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THE  NORTH  AMERICAN  SPECIES  OF 
THE  BEE  GENUS  UTHURGE 
(HYMENOPTERA:  MEGACHILIDAE) 


Roy  R.  Snelling 


Number  343 
5 July  1983 


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THE  NORTH  AMERICAN  SPECIES  OF 
THE  BEE  GENUS  LITHURGE 
(HYMENOPTERA:  MEGACHILIDAE) 

Roy  R.  Snelling1 


ABSTRACT.  The  North  American  species  of  Lithurge  are  re- 
viewed, and  a key  is  given  for  the  separation  of  both  sexes  of  all 
species.  Supraclypeal  structures  of  the  females  are  illustrated,  as  are 
the  labra)  structures  of  the  males.  Of  the  previously  described  native 
forms,  L.  apicalis  var.  opuntiae  Cockerell  and  L.  arizonensis  Cock- 
erell are  synonyms  of  L.  apicalis  Cresson  (NEW  SYNONYMY);  L. 
apicalis  subsp.  lilloralis  Cockerell  is  a valid  species  and  a senior 
synonym  of  L.  bruesi  Mitchell  (NEW  SYNONYMY).  One  new 
species,  L.  listrota.  is  described  from  southern  California  desert  re- 
gions. All  native  American  species  belong  to  the  subgenus  Lithur- 
gopsis. 

Two  representatives  of  the  nominate  subgenus  occur  in  the  New 
World.  The  Palaearctic  species,  L.  chrysurus  (Fonscolombe).  is  ad- 
ventive  in  the  eastern  United  States.  A South  American  species,  L. 
huberi  Ducke,  belongs  to  the  Indo-Australian  species  group  of  L. 
atratus  F.  Smith  and  may  have  been  introduced  into  Brazil  in  historic 
times. 

The  South  American  genera  Trichothurgus  Moure  and  Lithur- 
gomma  Moure  are  merged  into  a single  genus,  Trichothurgus,  and 
the  tribe  Trichothurgini  is  dropped. 

INTRODUCTION 

The  megachilid  subfamily  Lithurginae  includes  about  50 
species  in  both  Old  and  New  World  faunas.  Old  World  species 
all  belong  to  the  nominate  subgenus  of  Lithurge  Latreille  and 
are  present  in  the  southern  Palaearctic,  Ethiopian  and  Indo- 
Australian  regions.  These  species,  as  far  as  known,  provision 
their  nests  with  pollen  from  plants  of  the  families  Malvaceae 
and  Compositae. 

For  the  most  part.  New  World  species  of  Lithurge  belong 
to  the  subgenus  Lithurgopsis  Fox  and  are  present  in  southern 
North  America  and  South  America.  One  Palaearctic  species, 
L.  chrysurus  (Fonscolombe),  is  adventive  in  the  eastern  United 
States  (Roberts,  1978);  a Brazilian  species,  L.  huberi  Ducke, 
also  belongs  to  the  nominate  subgenus  and  is  probably  in- 
troduced from  the  Indo-Australian  region.  Although  most 
Lithurgopsis  utilize  pollen  from  Cactaceae  in  their  larval  cells, 
a few  Argentinian  species  use  malvaceous  pollens  (J.  L.  Neff, 
in  litt.).  Some  Chilean  and  Argentinian  species,  formerly 
placed  in  Lithurge,  are  now  placed  in  the  genera  Tricho- 
thurgus and  Lithurgomma,  both  described  by  Moure  (1949); 

Contributions  in  Science,  Number  343,  pp.  1-11 

Natural  History  Museum  of  Los  Angeles  County,  1983 


at  least  one  of  these  is  an  oligolege  on  Compositae  (J.  L.  Nelf, 
in  litt.). 

THE  NEW  WORLD  LITHURGINAE 

Most  New  World  Lithurginae  have  been  described  in  the 
genus  Lithurgus  Berthold,  1827,  an  invalid  emendation  of 
Lithurge  Latreille,  1825.  Fox  (1902)  noted  that  males  of  the 
New  World  species  possess  tarsal  arolia  and  proposed  to 
remove  them  to  a new  genus,  Lithurgopsis.  Subsequent  work- 
ers have  consistently  treated  Lithurgopsis  as  a subgenus  of 
Lithurge  (e.g.,  Michener,  1944;  Hurd,  1979). 

In  addition  to  the  presence  of  tarsal  arolia  in  Lithurgopsis 
males,  there  are  other  features  by  which  members  of  the 
subgenus  may  be  separated  from  the  nominotypical  subge- 
nus. In  females  of  Lithurgopsis  the  facial  prominence,  when 
present,  occurs  only  on  the  supraclypeal  area;  in  Lithurge, 
sensu  stricto,  the  basal  portion  of  the  clypeus  is  also  involved. 
The  first  flagellar  segment  is  consistently  short  in  both  sexes 
of  Lithurgopsis ; it  is  little,  if  any,  longer  than  the  second  and 
often  much  shorter;  the  second  segment  is  as  long  as  broad 
or  longer.  In  Lithurge,  sensu  stricto,  the  first  flagellar  segment 
is  as  long  as  broad,  or  longer,  the  second  segment  not  more 
than  half  as  long  as  the  first;  the  second  segment  is  often 
conspicuously  broader  than  long. 

Lithurge  (L.)  chrysurus  (Fonscolombe,  1834)  is  a Palaearc- 
tic species  that  is  now  introduced  and  established  in  the 
eastern  United  States.  Roberts  (1978)  reported  this  Medi- 
terranean species  from  New  Jersey,  discussed  its  nesting  bi- 
ology and  behavior,  and  described  the  larva  and  pupa. 

Lithurge  (L.)  huberi  (Ducke,  1907),  described  from  Brazil, 
is  the  only  South  American  representative  of  the  nomino- 
typical subgenus.  I have  seen  specimens  of  both  sexes  of  this 
species.  Morphologically,  L.  huberi  is  a member  of  the  group 
of  forms  related  to  L.  atratus  (F.  Smith).  Nominate  species 
of  this  group  range  from  India  to  Australia,  Melanesia,  and 
Polynesia;  according  to  Michener  ( 1965)  there  may  be  but  a 


1.  Entomology  Section,  Natural  History  Museum  of  Los  Angeles 
County. 


ISSN  0459-8113 


single  species  involved.  There  are  no  obvious  morphological 
features  by  which  L.  huberi  may  be  separated  from  L.  alratus. 
It  seems  likely  that  L.  huberi  is  adventive  in  Brazil,  intro- 
duced via  infested  wood  within  historic  times. 

Although  most  New  World  Lithurginae  belong  to  the  genus 
Litharge,  Moure  ( 1 949)  described  two  genera,  Trichothurgus 
and  Lithurgomma,  which  he  placed  in  the  new  tribe  Tricho- 
thurgini.  Species  of  Trichothurgini  occur  in  Peru  and  Argen- 
tina but  are  primarily  Chilean.  Trichothurgus  and  Lithur- 
gomma differ  from  Litharge  in  the  elongate  labrum  and  in 
the  shape  of  the  first  gastric  tergum,  which  is  not  flattened 
as  in  Litharge.  The  species  of  these  two  genera  are  large, 
robust  bees  with  abundant  body  hairs  that  conceal  most  in- 
tegumentary surfaces. 

In  Trichothurgus,  the  facial  prominence,  if  present,  is  on 
the  supraclypeal  area  and  the  frons  lacks  a shiny  area  in  front 
of  the  anterior  ocellus.  Species  of  Lithurgomma  have  a clyp- 
eal  prominence,  in  females  only,  and  there  is  a large,  flat- 
tened, shiny  area  in  front  of  the  anterior  ocellus.  These  dif- 
ferences are  trivial  and  do  not,  in  my  opinion,  justify  the 
status  of  Trichothurgus  and  Lithurgomma  as  separate  genera. 
My  view  is  that  Lithurgomma  is  a junior  synonym  of  Tricho- 
thurgus (NEW  SYNONYMY).  The  characters  of  Moure’s 
Trichothurgini  will  differentiate  the  expanded  genus  Tricho- 
thurgus from  Litharge.  Concurrently,  the  tribe  Trichothur- 
gini is  superfluous;  there  is  no  need  to  place  the  two  genera 
of  Lithurginae  in  separate  tribes. 

The  known  species  of  Trichothurgus  are:  T.  a/biceps  (Friese), 
T.  alpestris  (Friese),  T.  aterrimus  (Cockerell),  T.  colloncu- 
rensis  Ogloblin,  T.  herbsti  (Friese),  T.  holomelan  (Moure), 
T.  laticeps  (Friese),  T.  neoqueensis  (Friese),  T.  osmioides 
(Friese),  T.  pseudocellalus  (Moure),  T.  shajovskoyi  Ogloblin, 
and  T.  wagenknechti  (Moure). 

The  few  South  American  species  of  Lithurge  have  never 
been  critically  examined.  The  North  American  species  were 
reviewed  by  Mitchell  (1938),  who  gave  a key  for  the  species 
then  known.  The  biology  of  no  American  species  has  been 
studied  in  detail;  some  notes  on  L.  apicalis  (Cresson)  were 
published  by  Parker  and  Potter  (1973),  and  Brach  (1979) 
reported  some  observations  on  L.  gibbosa  (F.  Smith).  The 
nesting  biology  and  behavior  of  the  introduced  species,  L. 
chrysurus,  were  described  by  Roberts  (1978). 

SPECIMENS  EXAMINED 

During  the  course  of  this  study,  specimens  from  the  following 
institutional  collections  were  examined:  American  Museum 
of  Natural  Flistory  (AMNH),  Arizona  State  University 
(ARSU),  British  Museum  (Natural  History)  (BMNH),  Cal- 
ifornia Academy  of  Sciences  (CAS),  California  Department 
of  Food  and  Agriculture  (CDFA),  National  Museum  of  Nat- 
ural History  (USNM),  Natural  History  Museum  of  Los  An- 
geles County  (LACM),  San  Diego  Natural  History  Museum 
(SDM),  Texas  A and  I University  (TAI),  Texas  A and  M 
University  (TAMU),  University  of  Arizona  (UNAR),  Uni- 
versity of  California,  Berkeley  (UCB),  University  of  Califor- 
nia, Davis  (UCD),  University  of  California,  Riverside  (UCR), 
University  of  Colorado  (UCOL),  University  of  Kansas 


(UKAN),  University  of  Nebraska  (UNEB),  and  Utah  State 
University  (UTSU).  Specimens  from  the  personal  collection 
of  T.  Griswold  are  indicated  (GRIS). 

SYSTEMATICS 

In  the  review  by  Mitchell  (1938),  five  species  were  treated 
in  the  North  American  fauna,  one  with  three  subspecies.  A 
sixth  species  was  mentioned  but  not  treated  as  it  had  been 
too  recently  described  and  was  unknown  to  Mitchell.  The 
present  study  recognizes  eight  species,  including  one  adven- 
tive species  and  one  that  was  previously  undescribed.  The 
various  subspecies  attributed  to  L.  apicalis  by  Mitchell  ( 1 938) 
are  discussed  below.  The  morphological  characteristics  of 
each  species  are  sufficiently  distinctive  that  the  previously 
described  species  are  not  redescribed;  the  keys  and  figures 
given  here  are  adequate  for  their  recognition. 

Superficially,  the  various  species  tend  to  be  quite  similar. 
Females  are  most  notably  different  from  one  another  in  the 
shape  of  the  elevations  or  processes  of  the  supraclypeal  area. 
There  are  differences,  also,  in  the  density  of  the  punctures  of 
the  clypeus  and  supraclypeal  area,  but  there  is  sufficient  infra- 
specific variability  to  render  these  unreliable  as  diagnostic 
characters. 

Males  are  most  readily  separated  by  differences  in  labral 
structure.  When  fresh,  individuals  of  this  sex  should  have 
the  mandibles  spread  so  that  the  external  surface  of  the  la- 
brum is  clearly  visible.  Although  the  mandibles  of  a dry 
specimen  can  be  spread  after  relaxing  the  bee,  there  is  a 
greater  chance  that  the  specimen  will  be  damaged. 

KEY  TO  NORTH  AMERICAN  LITHURGE 
Males 

la.  Tarsal  arolia  present;  first  flagellar  segment  shorter  than 

second  (subgenus  Lithurgopsis) 2 

b.  Tarsal  arolia  absent;  first  flagellar  segment  distinctly 

longer  than  second  (subgenus  Lithurge)  

chrysurus  (Fonscolombe) 

2a.  Labrum  with  a single  median  tubercle,  or  none  ....  3 
b.  Labrum  with  a pair  of  erect,  slender  tubercles  (Fig.  4) 

littoralis  (Cockerell) 

3a.  Labrum  without  median  tubercle,  with  a low,  transverse 


basal  ridge  or  swelling  (Figs.  6,  10)  4 

b.  Labrum  with  an  erect,  median  tubercle,  without  trans- 
verse basal  ridge  or  swelling  (Figs.  8,  12)  6 


4a.  Supraclypeal  area  gently  convex,  its  upper  portion  slop- 
ing toward  antennal  sockets,  and  with  a definite  median 
area  that  is  less  closely  punctate  than  area  near  suban- 
tennal  suture  or  entire  supraclypeal  area  sparsely  and 

irregularly  punctate  5 

b.  Supraclypeal  area  uniformly  flat  between  clypeal  base 
and  antennal  sockets,  continguously  punctate,  punctures 

coarse  and  subcontiguous  along  midline 

planifrons  (Friese) 

5a.  First  flagellar  segment,  on  shortest  side,  distinctly  broad- 
er than  long,  shorter  than  pedicel;  supraclypeal  area  about 


2 Contributions  in  Science,  Number  343 


Snelling:  North  American  Lithurge 


1.4  times  wider  than  long;  legs  usually  reddish  

echinocacti  (Cockerell) 

b.  First  flagellar  segment,  on  shortest  side,  about  as  long  as 
wide,  about  as  long  as  pedicel;  supraclypeal  area  about 

1.8  times  wider  than  long;  legs  brown  

apicalis  (Cresson) 

6a.  Labrum  with  a low,  often  obsolescent,  ridge  extending 
obliquely  from  basal  corner  to  base  of  median  tubercle 

(Figs.  2,  12)  7 

b.  Labrum  deeply  concave  on  either  side  of  tubercle  and 
with  a short,  curved  ridge  from  base  of  tubercle  to  mid- 
length of  lateral  margin  (Fig.  8)  

socorroensis  (Mitchell) 

7a.  Supraclypeal  area  slightly  protuberant,  densely  punctate; 
ocelloccipital  distance  greater  than  interocellar  distance 

gibbosa  (F.  Smith) 

b.  Supraclypeal  area  flat,  sparsely  punctate,  at  least  in  mid- 
dle; ocelloccipital  distance  less  than  interocellar  distance 
listrota  new  species 

Females 

la.  First  flagellar  segment  shorter  than  second;  facial  prom- 
inence, when  present,  not  as  below  (subgenus  Lithur- 

gopsis) 2 

b.  First  flagellar  segment  distinctly  longer  than  second;  fa- 
cial prominence  high,  triangular  in  lateral  view,  its  lower 
portion  sloping  toward  base  of  clypeus  (subgenus  Li- 
tharge)   chrysurus  (Fonscolombe) 

2a.  Supraclypeal  protuberance  nearly  as  broad  as  face,  ex- 
tending laterad  of  subantennal  sutures  (Figs.  3,  5);  malar 
area  distinct  and  with  deep  posterior  pit  (Figs.  3,  5)  3 

b.  Supraclypeal  protuberance,  when  present,  much  narrow- 
er than  width  of  face,  not  extending  laterad  of  suban- 
tennal sutures  (Figs.  7,  9,  1 1 );  malar  area  linear,  mandible 
nearly  contiguous  with  eye,  posterior  pit  usually  absent 

(Figs.  1 , 7,  9) 4 

3a.  Labral  tubercle  with  median  and  sublateral  impressions; 

clypeus  without  preapical  ridge  (Fig.  3)  

/ itt oralis  (Cockerell) 

b.  Labral  tubercle  divided  in  middle  only  (Fig.  5);  clypeus 
with  preapical  ridge  which  is  interrupted  in  middle  (Fig. 

5)  apicalis  (Cresson) 

4a.  Supraclypeal  area  distinctly  elevated  on  each  side  of  mid- 
dle or  with  a single,  transverse,  bowed  elevation  .5 
b.  Supraclypeal  area  flat,  with  no  protuberance,  shiny  and 
very  sparsely  punctate  (Fig.  1)  ...  listrota  new  species 
5a.  Supraclypeal  area  with  prominent  rounded  elevation  on 
either  side  of  middle  (Figs.  7,  9);  malar  area  without 

posterior  pit  6 

b.  Supraclypeal  elevation  a high,  transverse,  bowed  ridge 
(Fig.  1 1 );  malar  area  with  broad,  shallow,  posterior  pit 

gibbosa  (F.  Smith) 

6a.  Dorsal  margin  of  pronotum  with  conspicuous  fascia  of 
dense,  plumose,  white  hairs;  ocelloccipital  distance  less 

than  interocellar  distance  echinocacti  (Cockerell) 

b.  Dorsal  margin  of  pronotum  without  fascia  of  dense,  plu- 


mose, white  hairs;  ocelloccipital  distance  greater  than 
interocellar  distance socorroensis  (Mitchell) 

Lithurge  ( Lithurgopsis ) apicalis  (Cresson) 

Figures  5,  6 

Lithurgus  apicalis  Cresson,  1875:724.  9. 

Lithurgopsis  apicalis  var.  opuntiae  Cockerell,  1 902;  182.9. 
NEW  SYNONYMY. 

Lithurgus  arizonensis  Cockerell,  1937:108.  9.  NEW  SYN- 
ONYMY. 

RANGE 

South  Dakota  and  Wyoming  to  western  Texas  (?),  west  to 
Utah,  Nevada,  and  southern  California;  northern  Mexico. 

Although  this  species  has  been  recorded  from  Texas  by 
previous  authors  (Cockerell,  1911;  Mitchell,  1938),  I have 
seen  no  material  of  L.  apicalis  from  that  State.  All  specimens 
from  Texas  previously  identified  as  L.  apicalis,  that  I have 
examined,  proved  to  be  L.  littora/is. 

BIONOMICS 

Parker  and  Potter  (1973)  published  some  notes  on  the  nesting 
habits  of  L.  apicalis.  The  larva  has  been  described  by  Rozen 
(1973). 

Females  provision  the  cells  with  pollen  from  species  of 
Opuntia,  but  both  sexes  have  been  taken  at  flowers  of  other 
genera.  Hurd  ( 1979)  recorded  this  species  on  flowers  of  Echi- 
nocactus,  Ence/ia  californica,  Eriogonum,  and  Sphaeralcea. 
Previously  unpublished  floral  records  include:  Agave  neva- 
densis,  Argemone  platyceros,  Argemone  sp.,  Asclepias  erosa, 
Baileya  multiradiata,  Cirsium  californicum,  Cirsium  sp., 
Calochortus  concolor,  Chi/opsis  linearis,  Cleome  serrulata, 
Cnicus  sp.,  Grindelia  squarrosa,  Prosopis  juliflora,  Robinia 
neomexicana,  Senecio  longi/obus,  Verbena  stricta,  Verbesina 
encelioides,  and  “sweet  pea.” 

DISCUSSION 

For  many  years,  L.  apicalis  has  been  divided  into  two  sub- 
species: L.  a.  apicalis  and  L.  a.  opuntiae.  The  former,  with 
ferruginous  hairs  on  the  last  tergum  of  the  female,  is  a north- 
ern form,  found  in  South  Dakota,  Wyoming,  Nebraska,  Kan- 
sas, Colorado,  Utah,  and  Nevada.  The  southern  population, 
L.  a.  opuntiae,  with  dark  brown  hairs  on  the  last  tergum  of 
the  female,  ranges  from  New  Mexico  to  southern  California. 
Both  forms,  however,  occur  in  New  Mexico,  Arizona,  and 
California.  In  many  areas  the  two  occur  together,  along  with 
many  specimens  of  intermediate  character. 

In  view  of  the  broad  zone  of  mtergradalion,  were  mixing 
of  the  forms  occurs  and  which  actually  includes  much  of  the 
range  of  L.  a.  opuntiae,  it  hardly  seems  desirable  to  separate 
the  two  forms  nomenclatorially.  Accordingly,  L.  a.  opuntiae 
its  here  treated  as  a synonym  of  L.  apicalis. 

Cockerell  (1937)  described  L.  arizonensis  from  two  fe- 
males collected  in  the  Baboquivari  Mountains  of  Arizona. 
These  were  said  to  differ  from  L.  a.  opuntiae  by  their  more 


Contributions  in  Science,  Number  343 


Snelling:  North  American  Lithurge  3 


Figures  1-6.  Lithurge  spp.  Figs.  1,  3,  5,  lower  portion  of  face  of  female  of  L.  listrota  (1),  L.  littoralis  (3).  and  L.  apicalis  (5).  Scale  line  = 1.0 
mm.  Figs.  2,  4,  6,  labrum  of  male  of  L.  listrota  (2),  L.  littoralis  (4),  and  L.  apicalis  (6).  Scale  line  = 0.5  mm. 


4 Contributions  in  Science,  Number  343 


Snelling:  North  American  Lithurge 


flattened  and  coarsely  punctate  clypeus  and  by  the  deeper, 
sparser  facial  punctures.  I have  examined  the  type  of  L. 
arizonensis  (in  the  CAS)  and  find  no  significant  differences 
between  this  specimen  and  other  individuals  of  L.  apicalis. 
All  the  differences  cited  by  Cockerell  fall  well  within  the  range 
of  character  variation  that  1 attribute  to  this  species. 

The  male  from  Port  Isabel,  Texas,  described  by  Cockerell 
( 1 9 1 7)  as  L.  apicalis  subsp.  littora/is,  is  a distinct  species  and 
a senior  synonym  of  L.  bruesi  (see  below). 

TYPE  MATERIAL 

Lithurgus  apicalis  Cresson:  female,  in  Academy  of  Natural 
Sciences,  Philadelphia.  Lithurgopsis  apicalis  var.  opuntiae 
Cockerell:  no  types  designated,  and  none  of  the  original  ma- 
terial can  be  identified  with  surety.  Lithurgus  arizonensis 
Cockerell:  female,  in  California  Academy  of  Sciences,  San 
Francisco. 

SPECIMENS  EXAMINED  (566  29,  423  33) 

UNITED  STATES.  ARIZONA,  Apache  Co.:  White  Mts. 
(CAS);  Carrizo  (UNAR).  Cochise  Co.:  Chiricahua  Mts.  (Bar- 
foot  Park,  8600  ft.  elev.;  Cave  Creek;  Pinery  Canyon;  Portal 
and  vicinity;  Southwest  Research  Station;  Sunny  Flat;  AMNFI, 
ARSU,  LACM,  UNAR,  UCB,  UCD,  UKAN,  UTSU);  Hua- 
chuca  Mts.  (Ramsey  Canyon;  UKAN,  UNAR);  Mustang  Mts. 
(UKAN);  6 mi.  SE  of  Willcox  (AMNH);  Douglas  (UCB). 
Coconino  Co.:  24  mi.  W of  Cameron  (UKAN,  UNEB);  Fre- 
donia  (UTSU);  Grand  Canyon  Natl.  Park  (LACM,  UCD, 
UNAR);  Flagstaff  and  vicinity  (UCB,  UNAR,  USNM);  Oak 
Creek  Canyon  (CAS,  LACM,  UKAN,  UNEB,  UTSU);  Se- 
dona  (UKAN,  UNAR).  Gila  Co.:  Florence  (ARSU);  18  mi. 
SE  of  Globe  (UKAN);  18  mi.  S of  Payson  (UNAR);  San 
Carlos  Lake  (UNAR).  Greenlee  Co.:  23  mi.  N of  Clifton 
(ARSU).  Maricopa  Co.:  Granite  Reef  Dam  (ARSU).  Navajo 
Co.:  Fort  Apache  (UNAR);  Cedar  Creek,  15  mi.  W of  Fort 
Apache  (UNAR);  16  mi.  SW  of  Kayenta  (ARSU);  Marsh 
Pass  (USNM,  UTSU).  Pima  Co.:  Baboquivari  Mts.  (Brown 
Canyon;  Elkhorn  Ranch  Canyon;  AMNH,  CAS,  LACM, 
UKAN,  LINAR);  Redington  and  Redington  Pass  (LINAR); 
Rincon  Mts.  (Madrona  Ranger  Station;  1.5  mi.  NE  of  X-9 
Ranch;  UNAR);  Santa  Rita  Mts.  (Gardner  Canyon;  Madera 
Canyon;  LACM,  UKAN,  UNAR);  Santa  Catalina  Mts.  (Sa- 
bino  Canyon;  UNAR);  Silver  Bell  bajada  (LACM);  Tucson 
and  vicinity  (UCD,  UNAR,  UNEB,  USNM,  UTSU).  Pinal 
Co.:  Oracle  (USNM);  Superior  (B.  Thompson  Arboretum; 
UNAR).  Santa  Cruz  Co.:  Atascosa  Mts.  (Sycamore  Canyon; 
UNAR);  Nogales  (USNM);  Patagonia  (UNAR);  Sonoita 
(LACM).  Yavapai  Co.:  Ash  Fork  (USNM);  Hell  Canyon 
(ARSU);  Jerome  and  vicinity  (UKAN,  UNEB);  Mingus  Mt. 
(UTSU);  5 mi.  NE  of  Paulden  (ARSU);  Prescott  and  vicinity 
(LACM,  UCB);  24  mi.  SE  of  Wikieup  (ARSU).  County  un- 
known: White  River  (UNAR;  this  locality  name  occurs  in 
three  different  counties).  CALIFORNIA,  Inyo  Co.:  Bristle- 
cone  Natl.  Forest  (Schulman  Grove;  UCD,  UTSU);  Argus 
Mts.  (LACM);  Mountain  Spring  (UCB);  Panamint  Mts.  (Ben- 
nett Peak,  9980  ft.  elev.;  Frenchman’s  Canyon,  6800-7200 
ft.  elev.;  GRIS,  UCR,  UTSU);  White  Mts.  (Wyman  Canyon. 
8500  ft.  elev.;  UCB).  Los  Angeles  Co.:  Eagle  Rock  (UKAN). 


Mono  Co.:  Benton  (UCD);  Cottonwood  Creek  (9300  ft.  elev.; 
UCB);  Mammoth  Lake  (UCD,  LITSU).  Riverside  Co.:  Anza 
and  vicinity  (CDFA,  LACM,  UCB,  UCD,  UTSU);  Banning 
(CAS);  Deep  Canyon  (UCB,  UCR);  Joshua  Tree  Natl.  Mon- 
ument (UCR);  Mt.  San  Jacinto  (UTSU);  4 mi.  S of  Palm 
Desert  (UCB);  Palm  Springs  (UCD);  Pinon  Flat  (LACM, 
UCB,  UKAN);  Ribbonwood  (UCB);  Santa  Rosa  Peak  (8000 
ft.  elev.;  CAS).  San  Bernardino  Co.:  Caruthers  Canyon  (5650 
ft.  elev.;  GRIS);  Cedar  Canyon  (5100  ft.  elev.;  GRIS);  9.5 
mi.  NNE  of  Cima  (4200-4265  ft.  elev.;  LACM);  Cima  Dome 
(5050  ft.  elev.;  GRIS);  Clark  Mtn.  (6000  ft.  elev.;  GRIS); 
Deep  Creek  Public  Camp  (LACM);  Dove  Spring  (4600  ft. 
elev.;  GRIS);  Falls  Public  Camp  (UCB);  Forest  Home  (CAS, 
UCR);  Granite  Mts.  (4050  ft.  elev.,  GRIS);  Joshua  Tree 
(LACM);  Lower  Covington  Flat  (LACM);  Mitchell’s  Caverns 
(GRIS);  s.  fork,  Santa  Ana  River  (UNEB);  Wheaton  Springs 
(UCB).  San  Diego  Co.:  Anza-Borrego  Desert  State  Park 
(LACM);  5 mi.  W of  Borrego  Springs  (UCD);  Jacumba 
(LACM);  10-1  1 mi.  E of  Julian  (LACM);  Laguna  (SDM); 
Mt.  Laguna  (UCB);  Oak  Grove  (UCR);  San  Felipe  Wash 
(LACM,  UCR);  2 mi.  N of  Warner  Springs  (LACM,  UCB, 
UCD,  UTSU).  COLORADO,  Alamosa  Co.:  Great  Sand 
Dunes  Natl.  Monument  (LACM,  UCOL,  UKAN).  Archuleta 
Co.:  Arboles  (USNM);  Yellow  Jacket  (UCOL).  Boulder  Co.: 
Boulder  and  vicinity  (AMNH,  UCOL,  UKAN);  Gregory 
Canyon  (AMNH);  Pinecliffe  (UKAN).  Chaffee  Co.:  Buena 
Vista  (7900-8000  ft.  elev.;  AMNH).  Costilla  Co.:  Fort  Gar- 
land (UNEB).  Crowley  Co.:  Fowler  (UCOL).  Delta  Co.:  5 
mi.  N of  Cedaredge  (UCOL);  Delta  (UCOL);  Paonia  (CAS). 
£7  Paso  Co.:  Manitou  (UCB,  UKAN,  USNM);  Ute  Creek 
(LJNEB).  Fremont  Co.:  Canon  City  (UKAN);  10  mi.  NE  of 
Cotopaxi  (5900  ft.  elev.;  UKAN).  Garfield  Co.:  Glenwood 
Springs  (AMNH).  Huerfano  Co.:  Cuchara  Dam  (UCOL); 
Lathrop  State  Park  (UCOL);  2-3  mi.  W of  Walsenberg 
(UCOL,  UKAN).  Mesa  Co.:  Colorado  Natl.  Monument 
(UCOL).  Moffat  Co.:  Dinosaur  Natl.  Monument  (Castle  Rock; 
Red  Rock  Ranch;  UCOL).  Montrose  Co.:  Naturita  (UKAN). 
Ouray  Co.:  Ridgway  (AMNH).  KANSAS,  Hamilton  Co.: 
(UKAN).  Kingman  Co.:  Calista  (UKAN).  Pratt  Co.:  8 mi. 
NE  of  Isabel  (UKAN).  NEBRASKA,  Arthur  Co.:  4 mi.  N of 
Arthur  (UKAN).  Sioux  Co.:  Warbonnet  Canyon  (UNEB). 
NEVADA,  Clark  Co.:  (SDM);  Charleston  Mts.  (Lee  Canyon; 
Willow  Creek  Camp.  AMNH,  UCD).  Humboldt  Co.:  Win- 
nemucca  (UTSU).  NEW  MEXICO,  Bernalillo  Co.:  Cedro 
Canyon  (UKAN);  Cienega  Canyon  (UKAN);  Sandia  Mts. 
(UKAN).  Chavez  Co.:  Elk  (UNAR).  Dona  Ana  Co.:  Las 
Cruces  (AMNH,  BMNH,  UCOL,  UKAN,  UTSU).  Grant 
Co.:  13  mi.  N of  Silver  City  (6900  ft.  elev.;  AMNH).  Gua- 
dalupe Co.:  Vaughn  (UNEB).  Hidalgo  Co.:  Post  Office  Can- 
yon (ARSU).  Lincoln  Co.:  Alto  (UKAN);  Capitan  (UNEB); 
Lincoln  (LJKAN);  Nogal  (UKAN,  UNEB);  Ruidoso  and  vi- 
cinity (AMNH,  UKAN).  McKinley  Co.:  Pinedale  (USNM). 
Otero  Co.:  High  Rolls  (AMNH).  Rio  Arriba  Co.:  Ghost  Ranch 
(UNAR);  Echo  Canyon  (UCR).  Sandoval  Co.:  Jemez  Spring 
(AMNH);  10  mi.  N of  Jemez  Spring  (7500  ft.  elev.;  AMNH); 
San  Ysidro  (AMNH).  San  Juan  Co.:  Bloomfield  (UTSU). 
Santa  Fe  Co.:  Lamy  (USNM);  Nambe  (UTSU);  Santa  Fe 
and  vicinity  (CAS,  UKAN,  UNEB).  Torrance  Co.:  Durran 


Contributions  in  Science,  Number  343 


Snelling:  North  American  Lithurge  5 


(UCB).  OKLAHOMA,  Custer  Co.:  Weatherford  (UKAN). 
Cimarron  Co.:  Black  Mesa  (UTSU).  SOUTH  DAKOTA, 
Custer  Co.:  Custer  (UNEB).  UTAH,  Beaver  Co.:  Beaver 
(UTSU);  16  mi.  E of  Beaver  (Keat’s  Lake,  8800  ft.  elev.; 
AMNH).  Cache  Co.:  Logan  (UCD).  Duchesne  Co.:  Roosevelt 
(UTSU).  Garfield  Co.:  Escalante  River  (UTSU).  Grand  Co.: 
Castle  Valley  (UTSU).  Juab  Co.:  Eureka  (UTSU).  Kane  Co.: 
Kanab  (UTSU);  Navajo  Mt.  (UTSU).  San  Juan  Co.:  5 mi. 
W of  Monticello  (Dalton  Springs,  850  ft.  elev.;  AMNH). 
Tooele  Co.:  (UCD).  Uintah  Co.:  Vernal  (UTSU).  Utah  Co.: 
Thistle  (UTSU).  WYOMING,  Albany  Co.:  Laramie  and  vi- 
cinity (UKAN,  UTSU).  Sweetwater  Co.:  Green  River 
(AMNH).  Weston  Co.:  Newcastle  (USNM).  MEXICO.  BAJA 
CALIFORNIA:  Canon  del  Tajo  (LACM);  3 mi.  S of  Encinas 
(UCB)  SONORA:  San  Bernardo  (LACM). 

Lithurge  ( Lithurgopsis ) echinocacti  (Cockerell) 

Figures  9,  10 

Lithurgus  echinocacti  Cockerell,  1898:453.  9. 

RANGE 

New  Mexico  to  southern  California;  northwestern  Mexico, 
south  to  Nayarit. 

BIONOMICS 

In  Arizona,  L.  echinocacti  has  been  observed  nesting  in  rotted 
wood  (Parker  and  Potter,  1973).  Females  collected  near  Sa- 
huarita,  Pima  County,  Arizona,  are  noted  to  have  been  “bor- 
ing in  post  attached  to  house.” 

Known  pollen  sources  for  females  are  various  species  of 
Echinocactus.  Other  lloral  records  for  this  bee  include  Ar- 
gemone  sp.,  Baccharis  sp.,  Baileya  pleniradiata.  Chilopsis 
linearis.  Ferocactus  wis/izeni.  Heterotheca  sp.,  Parkinsonia 
aculeata.  and  Tamarix  pentandra. 

DISCUSSION 

This  appears  to  be  an  uncommon  species.  Females  may  be 
easily  recognized  by  the  combination  of  two  conical  protu- 
berances on  the  supraclypeal  area,  the  presence  of  a pubescent 
fascia  on  the  dorsal  margin  of  the  pronotum,  and  the  usually 
red  legs.  Males  are  similar  to  those  of  L.  apicalis  in  that  both 
lack  labral  tubercles,  but  differ  in  the  narrower  supraclypeal 
area. 

TYPE  MATERIAL 

Lithurgus  echinocacti  Cockerell:  female,  present  location  un- 
known. 

SPECIMENS  EXAMINED  (38  99,  97  <?<$) 

UNITED  STATES.  ARIZONA,  Cochise  Co.:  Apache  Pass 
(LACM);  12  mi.  E of  Benson  (Texas  Canyon,  4800  ft.  elev.; 
LACM);  Portal  and  vicinity  (LACM,  UCB,  UCR).  Graham 
Co.:  Dripping  Spring,  Whitlock  Mts.  (LINAR).  Pima  Co.: 
Florida  Wash  (SDM);  Santa  Rita  Mts.  (Box  Canyon;  Madera 


Canyon;  IBP  site;  AMNH,  LACM,  UCD,  UNAR,  UTSU); 
Silver  Bell  bajada  (LACM);  Mountain  View  (LACM,  UCR); 
Santa  Catalina  Mts.  (Sabino  Canyon;  Peppersauce  Canyon; 
AMNH,  UCR,  UNAR);  Continental  (UCR,  USNM);  Ba- 
boquivari  Mts.  (Brown  Canyon;  Kitts  Peak;  AMNH,  UKAN, 
UNAR);  Sahuarita  (UNAR);  Tucson  and  vicinity  (AMNH, 
CAS,  UCB,  UKAN,  UNAR,  UTSU).  Pinal  Co.:  Oracle  and 
vicinity  (CAS,  LACM,  UCR);  Ray  (ARSU);  Superior  (B. 
Thompson  Arboretum;  UNAR).  Santa  Cruz  Co.:  20  mi.  N 
of  Nogales  (Tumacacori;  UNAR);  Pena  Blanca  (UCD).  CAL- 
IFORNIA, Inyo  Co.:  5 mi.  E of  Valley  Wells  (UCR).  NEW 
MEXICO,  Hidalgo  Co.:  19  mi.  N of  Rodeo  (UTSU).  MEXI- 
CO. BAJA  CALIFORNIA:  13  mi.  SE  of  Millers  Landing 
(ARSU);  1 1 mi.  E of  Rosalillito  (UNAR);  13  km.  NW  of 
Rosarito  (LACM);  Valle  de  Santa  Maria  (LACM).  BAJA 
CALFORNIA  SUR:  48  km.  S of  Loreto  (LACM);  Los  Ba- 
rriles  (CDFA).  NAYARIT:  43  mi.  N of  Tepic  (UCB).  SO- 
NORA: Guaymas  (UTSU);  Rio  Mayo  (UTSU);  San  Bernar- 
do (LACM). 

Lithurge  ( Lithurgopsis ) gihbosa  (F.  Smith) 

Figures  11,  12,  13 

Lithurgus  gibbosus  F.  Smith,  1853:147.  9. 

Lithurgus  compressus  F.  Smith,  1853:147.  <3. 

RANGE 

North  Carolina  to  Florida,  west  to  Kansas,  Oklahoma,  and 
Texas. 

BIONOMICS 

Notes  on  the  nesting  biology  of  L.  gibbosa  were  published 
by  Brach  (1979).  Females  provision  cells  with  pollen  from 
species  of  Opuntia.  Mitchell  (1962)  has  recorded  this  bee 
from  flowers  of  the  genera  Cirsium.  Helianthus,  Ilex.  Pon- 
taderia,  and  Rudbeckia.  I have  also  seen  a few  males  taken 
at  Centaurea  americana.  Ratibida  sp.,  and  Teucrium  sp.  in 
Texas. 

DISCUSSION 

The  female  of  L.  gibbosa  is  easily  recognized  by  the  high, 
bowed  supraclypeal  process,  similar  to  that  of  members  of 
the  Palaearctic  species  allied  to  L.  cornuta  (Fabricius). 

Males  are  recognizable  by  the  combination  of  densely 
punctate  supraclypeal  area,  the  presence  of  a single  labral 
tubercle,  and  the  long  ocelloccipital  distance.  The  labral  tu- 
bercle is  a curved,  transverse,  subapical  ridge;  in  the  center, 
this  ridge  is  elevated.  Males  from  Florida  and  Georgia  have 
this  median  elevation  abrupt  and  quite  prominent  (Fig.  12). 
The  median  elevation  in  males  from  Texas  is  less  pronounced 
(Fig.  13).  This  difference  between  specimens  from  the  two 
areas  is  consistent  but  does  not  correlate  with  other  features. 

TYPE  MATERIAL 

Lithurgus  gibbosus  F.  Smith:  females,  British  Museum  (Nat- 


6 Contributions  in  Science,  Number  343 


Snelling:  North  American  Lithurge 


Figures  7-13.  Lithurge  spp.  Figs.  7,  9,  1 1,  lower  portion  of  face  of  female  of  L.  socorroensis  (7).  L.  echinocacti  (9),  and  L.  gibbosa  (11).  Scale 
line  = 1.0  mm.  Figs.  8,  10,  12,  13,  labrum  of  male  of  L.  socorroensis  (8),  L.  echinocacti  (10),  and  L.  gibbosa  from  Florida  and  Texas  (12  and 
13,  respectively).  Scale  line  = 0.5  mm. 


Contributions  in  Science,  Number  343 


Snelling:  North  American  Lithurge  7 


ural  History).  Lithurgus  compressus  F.  Smith:  male,  British 
Museum  (Natural  History). 

SPECIMENS  EXAMINED  (123  29,  98  33) 

UNITED  STATES  (western  records  only).  KANSAS,  King- 
man  Co.:  Calista  (UKAN).  OKLAHOMA,  Caddo  Co.:  Red 
Rock  Canyon  State  Park  (UKAN).  TEXAS.  Angelina  Co.: 
10  mi.  N of  Lufkin  (AMNH).  Bastrop  Co.:  Bastrop  and  vi- 
cinity (UKAN,  TAMU);  McDade  (UKAN).  Bexar  Co.: 
(TAMU,  UTSU);  San  Antonio  (USNM).  Brazos  Co.:  (TAMU, 
UCB);  College  Station  (TAMU).  Crockett  Co.:  (TAMU).  De 
Witt  Co.:  Cuero  (AMNH).  Gillespie  Co.:  Fredricksburg 
(USNM).  Goliad  Co.:  (TAMU).  Gonzales  Co.:  Palmetto  State 
Park  (UCB).  Howard  Co.:  Big  Spring  (USNM).  Jackson  Co.: 
Edna  (UKAN).  Kerr  Co.:  Kerrville  (LISNM).  La  Salle  Co.: 
Cotulla  (USNM).  Lee  Co.:  Fedor  (AMNH,  USNM);  Gid- 
dings  (UKAN).  Matagorda  Co.:  Matagorda  (UKAN).  Mav- 
erick Co.:  (AMNH).  Robertson  Co.:  (TAMU).  Shackleford 
Co.:  3 mi.  S of  Throckmorton  (UKAN).  Torrant  Co.:  Fort 
Worth  (TAMU).  Zava/la  Co.:  Nueces  River  (USNM). 

Lithurge  ( Lithurgopsis ) listrota,  new  species 

Figures  1,  2 

DIAGNOSIS 

Both  sexes  are  separable  from  all  previously  described  Li- 
thurgopsis by  the  flat  supraclypeal  area  with  a shiny  im- 
punctate  median  line  that  extends  onto  the  clypeus;  the  male 
is  further  distinguished  from  L.  planifrons,  the  only  other 
flat-faced  species,  by  possessing  a labial  tubercle. 

DESCRIPTION 

FEMALE.  Measurements  (mm).  Head  width  3.2,  head 
length  2.7,  wing  length  7.5,  total  length  12. 

Head.  About  1.2  times  broader  than  long;  eyes  moderately 
convergent  below,  upper  interorbital  distance  about  1 .2  times 
lower;  malar  space  linear,  without  posterior  pit.  Transverse 
basal  ridge  of  labrum  low,  broadly  interrupted  in  middle. 
Apical  margin  of  clypeus  not  thickened.  Supraclypeal  area 
flat,  about  1 .4  times  broader  than  long.  Clypeus  polished 
between  subcontiguous  to  dense  punctures;  supraclypeal  area 
broadly  impunctate  and  polished  in  middle,  elsewhere  sub- 
contiguously  punctate,  interspaces  polished.  Interocellar  dis- 
tance slightly  less  than  ocelloccipital  distance  or  ocellocular 
distance.  First  flagellar  segment  shorter  than  pedicel  or  sec- 
ond flagellar  segment;  median  flagellar  segments  distinctly 
broader  than  long. 

Thorax.  Anterior  half  of  mesoscutal  dorsum  sharply  sca- 
brous; posterior  half  of  mesoscutum  and  all  of  scutellum 
rugosopunctate. 

Color.  Blackish  brown;  legs  medium  brown;  tegula  and 
underside  of  flagellum  ferruginous. 

Pilosity.  Mostly  whitish;  reddish  hairs  along  apical  margin 
of  clypeus,  lower  margin  of  mandible,  and  inner  side  of  basi- 
tarsi;  first  five  gastric  terga  with  distinct  fasciae  of  dense, 
appressed  white  hairs;  posterior  margin  of  pronotum  with 


fascia  of  white,  plumose  hairs;  discs  of  terga  two  through  four 
with  sparse,  very  short,  fuscous  hairs;  fifth  segment  with 
conspicuously  longer,  denser  dark  hairs;  apical  segment 
densely  covered  with  long,  dark  brown  hairs;  scopa  whitish; 
all  sterna,  except  last,  with  conspicuous  apical  fascia  of  dense, 
appressed,  white  hairs,  last  segment  with  brown  hairs. 

MALE.  Measurements  (mm).  Head  width  2.5  to  3.2,  head 
length  2.0  to  2.6,  wing  length  5.7  to  7.5,  total  length  8.3  to 
10.5. 

Head.  Head  1 .2  to  1.3  times  broader  than  long;  eyes  mod- 
erately convergent  below,  upper  interorbital  distance  about 
1.2  times  lower  interorbital  distance.  Labrum  with  subbasal, 
median,  erect  tubercle  and  with  an  obsolete  ridge  extending 
from  base  of  tubercle  to  each  basal  corner.  Supraclypeal  area 
flat,  about  1 .4  times  broader  than  long.  Clypeus  weakly  shiny 
between  continguous  to  subcontiguous  punctures.  Supra- 
clypeal area  polished  and  nearly  impunctate  in  middle,  with 
subcontiguous  punctures  laterad  that  are  coarser  than  those 
of  clypeus.  Interocellar  distance  a little  greater  than  either 
ocelloccipital  distance  or  ocellocular  distance.  First  flagellar 
segment  about  as  long  as  pedicel,  a little  shorter  than  second 
flagellar  segment. 

Thorax.  Mesoscutum  and  scutellum  with  distinct,  contig- 
uous punctures. 

Color.  As  described  for  female. 

Pilosity.  Similar  to  that  of  female,  but  hairs  at  apex  of 
clypeus  whitish,  dark  hairs  of  discs  of  gastric  terga  longer; 
fasciae  of  second  and  third  terga  interrupted  in  middle;  apical 
fasciae  of  sterna  longer  than  in  female;  apical  sternite  with 
pale  hairs. 

TYPE  MATERIAL 

All  type  material  is  from  San  Bernardino  County,  California. 
Holotype  female:  Cedar  Canyon,  4650  ft.  elev.  (Sec.  36,  T13N, 
R14E),  18  June  1980  (T.  Griswold),  on  Eriogonum  fasci- 
culatum  ssp.  polifolium.  Allotype:  same  locality,  13  June 
1980  (T.  Griswold).  Paratypes:  2 22,  5 33,  same  locality,  18 
and  30  June  1980  (T.  Griswold);  1 9,  Fenner  Valley,  3670 
ft.  elev.  (Sec.  29,  T11N,  R15E).  17  June  1980  (T.  Griswold), 
on  Chilopsis  linearis ; 1 3,  Colton  Hills,  3500  ft.  elev.  (Sec. 
12,  T10N,  R14E),  17June  1 980  (T.  Griswold),  on  Ferocactus 
acanthodes ; 1 2,  7 33,  Wild  Horse  Canyon,  4300  ft.  elev.  (Sec. 
18,  T1  IN,  R15E),  6 June  1980  (T.  Griswold),  on  Opuntia 
echinocarpa ; 1 3,  same  locality,  27  May  1980  (T.  Griswold); 

1 2,  Van  Winkle  Mts.,  10  June  1965  (G.  E.  Wallace);  1 1 33, 
9.5  mi.  NNEofCima,  4200-4265  ft.  elev.  (Morningstar  Mine 
Road),  1-4  June  1977  (R.  R.  Snelling  and  C.  D.  George),  on 
Encelia  californica ; 1 3,  10  mi.  NE  of  Earp,  31  March  1959 
(A.  S.  Menke  and  L.  A.  Stange).  Holotype  and  allotype  in 
LACM;  paratypes  in  LACM,  UCR.  and  GRIS. 

ADDITIONAL  SPECIMENS 

CALIFORNIA,  Riverside  Co.:  1 2,  30  May  1963,  on  Opuntia 
ramosissima\  2 33,  3 May  1963,  on  Echinocactus  sp.;  1 3,  26 


8 Contributions  in  Science,  Number  343 


Snelling:  North  American  Lithurge 


April  1963,  on  Cereus  sp.;  all  from  Deep  Canyon  (E.  I.  Schlin- 
ger;  UCR). 

ETYMOLOGY 

From  Greek,  listrotos  (leveled),  in  reference  to  the  flat  su- 
praclypeal  area  of  both  sexes. 

BIONOMICS 

Little  is  known  of  this  species.  Floral  records  within  the 
Cactaceae  include  the  genera  Cereus,  Echinocactus,  Fero- 
cactus,  and  Opuntia  but  are  so  few  that  no  clear  preference 
can  be  established.  The  few  females  available  all  bear  pollen 
from  Cactaceae.  Noncactus  floral  visits  are  known  for  both 
sexes  and  are  presumed  nectar  sources. 

In  the  Mojave  Desert,  from  which  most  specimens  were 
collected,  the  elevational  range  is  from  3500  to  4650  feet 
(approximately  1067  to  1417  meters).  Many  specimens  do 
not  bear  elevation  data,  but  most,  if  not  all,  seem  to  fall 
within  this  range.  Most  specimens  have  been  collected  in 
June,  but  the  earliest  record  is  3 1 March.  Within  the  Mojave 
Desert,  captures  range  between  27  May  and  30  June.  Earlier 
records  are  all  from  the  margins  of  the  Colorado  Desert  (Deep 
Canyon  and  10  miles  northeast  of  Earp). 

DISCUSSION 

Both  sexes  of  L.  listrota  may  be  readily  separated  from  all 
known  species  of  Lithurgopsis  by  the  flattened  supraclypeal 
area  with  a shiny,  impunctate  median  line.  Since  the  females 
of  other  species  all  possess  some  conspicuous  modification 
of  the  supraclypeal  area,  the  lack  of  such  in  the  case  of  the 
L.  listrota  female  is  especially  obvious. 

Males  of  L.  listrota  differ  less  conspicuously  from  those  of 
the  other  species,  since  males  of  none  have  prominent  ele- 
vations on  the  supraclypeal  area.  But,  in  these  other  species, 
the  supraclypeal  area  is  weakly  depressed  along  the  midline 
and  weakly  elevated  toward  the  subantennal  suture  on  either 
side.  This  contour  is  particularly  evident  in  such  species  as 
L.  apicalis.  Only  in  L.  listrota  and  L.  planifrons  is  the  su- 
praclypeal area  completely  flat,  even  though  it  is  slightly 
elevated  above  the  adjacent  areas  of  the  face;  males  of  L. 
planifrons  have  the  median  line  coarsely  and  contiguously 
punctate  and  lack  an  erect  labial  tubercle. 

Of  the  known  North  American  species,  only  one,  L.  api- 
calis, is  sympatric  with  L.  listrota.  Males  of  L.  apicalis  tend 
to  be  larger  than  those  of  L.  listrota  and  possess  a relatively 
strongly  elevated  supraclypeal  area.  In  addition,  the  labrum 
does  not  bear  a median  tubercle  but  has  instead  a low  sub- 
basal  ridge. 

It  is  possible  that  the  ranges  of  L.  listrota  and  L.  echinocacti 
may  overlap  in  southern  California  or  in  Baja  California. 
The  male  of  L.  echinocacti  lacks  a median  tubercle  on  the 
labrum,  and  the  legs  are  often  ferruginous. 

Geographically,  the  nearest  species  with  a labial  tubercle 
is  L.  socorroensis  in  southern  Arizona.  In  this  species,  there 
is  a distinct  ridge  extending  laterad  from  the  median  tubercle 
to  the  margin  of  the  labrum  near  midlength.  In  L.  listrota 


males,  the  ridge  extends  obliquely  to  the  anterolateral  corner 
of  the  labrum.  In  addition,  the  clypeus  of  L.  socorroensis 
males  is  coarsely  punctate,  with  the  punctures  discrete,  es- 
pecially in  the  middle.  In  contrast,  the  clypeus  of  L.  listrota 
is  finely  and  contiguously  punctate  in  most  areas,  with  the 
punctures  often  distorted. 

Lithurge  ( Lithurgopsis ) littoralis  (Cockerell) 

Figures  3,  4 

Lithurgus  apicalis  littoralis  Cockerell,  1917:191.  3. 
Lithurgus  bruesi  Mitchell,  1927:104.  <3.  NEW  SYNONY- 
MY. 

RANGE 

Illinois  to  New  Mexico,  south  to  the  State  of  Oaxaca,  Mexico. 

BIONOMICS 

Parks  ( 1930)  observed  the  behavior  of  this  bee  (as  L.  bruesi) 
in  Texas.  He  noted  that  females  visited  Opuntia  lindheimeri, 
O.  ellisiana,  and  Echinocactus  setispinosus  for  pollen.  Nectar 
sources  for  both  sexes  included  Monarda  punctata,  M.  ci- 
triodora,  and  Vitex  negundo  incisa.  Previously  unpublished 
floral  records  include  Acacia  sp.,  Callirhoe  involucrata,  Col- 
ubrina  texensis,  Opuntia  macrorhiza,  and  “composite.” 

DISCUSSION 

The  type  of  L.  littoralis,  from  Port  Isabel,  Texas,  is  in  the 
British  Museum  (Natural  History)  and  was  made  available 
by  G.  R.  Else.  This  type  has  been  compared  with  specimens 
of  L.  bruesi ; there  is  no  doubt  that  the  two  are  conspecific. 
Mitchell  ( 1 938)  first  suggested  that  this  might  be  the  case  but 
had  not  seen  the  type  of  L.  littoralis. 

TYPE  MATERIAL 

Lithurgus  apicalis  littoralis  Cockerell:  male,  in  British  Mu- 
seum (Natural  History).  Lithurgus  bruesi  Mitchell:  male,  in 
Museum  of  Comparative  Zoology. 

SPECIMENS  EXAMINED  (86  2$,  87  33) 

UNITED  STATES.  ILLINOIS,  Jersey  Co.:  Principia  Col- 
lege, Elsah  (UKAN).  KANSAS,  Bourbon  Co.:  Fort  Scott 
(UKAN).  Johnson  Co.:  Olathe  (UKAN).  NEW  MEXICO, 
Lincoln  Co.:  Nogal  (UKAN).  OKLAHOMA,  Cimarron  Co.: 
14-18  mi.  N of  Boise  City  (UTSU).  TEXAS,  Aransas  Co.: 

1 0 mi.  N of  Rockport  (UKAN,  UNEB).  Bexar  Co.:  (AMNH. 
LACM,  TAMU,  UKAN,  UTSU);  San  Antonio  (TAMU);  Ft. 
Sam  Houston  (UCB).  Brewster  Co.:  Big  Bend  Natl.  Park  (The 
Basin;  Window  Trail;  AMNH,  TAMU,  UCB,  UKAN);  12 
mi.  NW  of  Alpine  (UKAN).  Cameron  Co.:  18  mi.  N of 
Harlingen  (UKAN);  McAllen  (AMNH);  Port  Isabel  (BMNH). 
Duval  Co.:  8 mi.  W of  Fremont  (TAI).  Goliad  Co.:  (TAMU). 
Hidalgo  Co.:  Edinburg  (AMNH).  Howard  Co.:  Big  Spring 
(USNM).  Jeff  Davis  Co.:  Davis  Mts.  (AMNH,  UKAN).  Jim 
Wells  Co.:  (TAMU).  Kenedy  Co.:  3 mi.  S of  Sarita  (TAI). 


Contributions  in  Science,  Number  343 


Snelling:  North  American  Lithurge  9 


Kerr  Co.:  Kerrville  (UKAN).  Kleberg  Co.:  Kingsville  and 
vicinity  (TAI,  UKAN);  Bishop  (UKAN).  La  Salle  Co.:  Co- 
tulla  (USNM).  Maverick  Co.:  Quemado  (UKAN).  Presidio 
Co.:  10  mi.  SE  of  Presidio  (TAMU).  San  Patricio  Co.: 
(UKAN).  Sutton  Co.:  Sonora  (UKAN);  45  mi.  S of  Sonora 
(UKAN).  Tom  Green  Co.:  (UKAN).  Travis  Co.:  Austin 
(TAMU).  Uvalde  Co.:  Garner  State  Park  (TAMU);  Sabinal 
(USNM);  Uvalde  (UKAN,  UNEB).  Val  Verde  Co.:  Del  Rio 
(AMNH);  1 9 mi.  S of  Langtry  (LACM).  Victoria  Co.:  Victoria 
(USNM).  Webb  Co.:  22  mi.  S of  Laredo  (UKAN).  Zavalla 
Co.:  Nueces  River  (USNM).  County  unknown:  “Lopeno” 
(UKAN).  MEXICO.  COAHUILA:  15  mi.  N of  Saltillo 
(AMNH).  HIDALGO:  Pachuca  (UTSU).  NUEVO  LEON:  4 
mi.  W of  El  Cercado  (UCB,  UKAN).  OAXACA:  Matatlan 
(UCB).  PUEBLA:  10.5  mi.  E of  Azumbilla  (TAMU). 

Litluirge  ( Lithurgopsis ) planifrons  (Friese) 

Litburgus  planifrons  Friese,  1908:62.  <3. 

RANGE 
Central  Mexico. 

DISCUSSION 

No  authentic  material  of  L.  planifrons  has  been  available  for 
study;  the  species  is  based  on  a unique  male,  and  the  original 
description  is  broad  enough  that  it  can  be  applied  to  almost 
any  North  American  Lithurge  male.  Were  it  not  for  the  de- 
scriptive specific  epithet  chosen  by  Friese,  this  would  remain 
a species  of  questionable  identity. 

Three  males  from  Chamela,  Jalisco,  Mexico,  collected  by 
S.  H.  Bullock,  are,  I believe,  representatives  of  this  species. 
More  important  than  that  they  agree  with  the  original  de- 
scription is  that  the  face  is  flat.  The  supraclypeal  area  is  not 
at  all  elevated  above  the  level  of  the  clypeus  and  does  not, 
in  its  upper  portion,  slope  toward  the  antennal  sockets.  The 
only  other  species  with  a flat  supraclypeal  area  is  L.  listrota ; 
males  of  L.  listrota  possess  a prominent  erect  tubercle  on  the 
labrum,  lacking  in  L.  planifrons.  In  addition,  males  of  L. 
listrota  have  a conspicuous  median  impunctate  line  on  the 
supraclypeal  area. 

Since  the  male  of  L.  planifrons  lacks  an  erect  labral  tu- 
bercle, it  resembles  males  of  both  L.  apica/is  and  L.  ecbino- 
cacti.  From  both  of  these  species,  L.  planifrons  is  separable 
by  the  flat  supraclypeal  area,  which  does  not  have  a median 
impunctate  line.  Instead,  the  supraclypeal  area  is  conspicu- 
ously more  coarsely  punctate  in  the  middle  than  on  either 
side;  these  coarse  punctures  are  subcontiguous,  and  a similar 
central  line  of  coarse  punctures  is  present  on  the  clypeus. 
The  supraclypeal  area  of  L.  apica/is  and  L.  echinocacti  is 
distinctly  convex  between  the  clypeal  base  and  the  antennal 
sockets  (often  depressed  in  the  center  in  L.  apica/is ),  and  the 
middle  is  distinctly  shiny  and  impunctate  (sometimes  the 
entire  supraclypeal  area  sparsely  punctate  in  L.  apicalis). 

The  type  locality  for  L.  planifrons  was  originally  cited  as 


“Jacubaya,  Mexico.”  This  is  a misprint  for  Tacabaya,  now 
a suburb  of  Mexico  City. 

Lithurge  ( Lithurgopsis ) socorroensis  (Mitchell) 

Figures  7,  8 

Litburgus  socorroensis  Mitchell,  1938:152-154.  2 <3. 
RANGE 

Southern  Arizona  to  Costa  Rica;  Revillagigedo  Archipelago. 

BIONOMICS 

Little  is  known  of  this  species,  not  previously  recorded  from 
the  mainland.  Both  sexes  have  been  taken  at  flowers  of  Opun- 
tia  sp.  on  Socorro  Island,  the  type  locality.  Males  were  taken 
in  Arizona  on  “thistle.”  In  Costa  Rica,  both  sexes  have  been 
collected  on  Opuntia  sp.  and  males  were  on  Cydista  heter- 
ophylla.  Paullinia  costaricensis,  Pterocarpus  rohrii,  Securi- 
daca  sylvestris,  and  “pink  bignon  vine.” 

DISCUSSION 

Females  of  this  species  are  similar  to  those  of  L.  echinocacti 
in  the  structure  of  the  supraclypeal  area.  In  L.  socorroensis, 
the  legs  are  brown,  rather  than  red,  and  there  is  no  pubescent 
fascia  on  the  dorsal  margin  of  the  pronotum.  The  male  can 
be  recognized  by  the  structure  of  the  labrum,  as  noted  in  the 
key. 

TYPE  MATERIAL 

Litburgus  socorroensis  Mitchell:  female  holotype  and  allo- 
type in  California  Academy  of  Sciences. 

SPECIMENS  EXAMINED  (18  22,  24  3<5) 

UNITED  STATES.  ARIZONA,  Pima  Co.:  Santa  Catalina 
Mts.  (UNAR);  Rincon  Mts.  (Madrona  Ranger  Station; 
UNAR);  Baboquivari  Mts.  (UCR);  Continental  (UTSU). 
MEXICO.  JALISCO:  Guadalajara  (UTSU).  MORELOS: 
Cuernavaca  (UKAN).  PUEBLA:  Petlalcingo  and  vicinity 
(UCD,  UTSU).  ISLAS  REVILLAGIGEDO:  Socorro  Island 
(CAS,  LACM,  UTSU).  SINALOA:  50  mi.  NW  of  Mazatlan 
(UTSU).  VERA  CRUZ:  8 mi.  SW  of  Vera  Cruz  (UTSU). 
EL  SALVADOR.  LA  LIBERTAD:  5 km.  NW  of Quezaltepec 
(Hacienda  Capolinas;  UCR).  COSTA  RICA.  GUANACAS- 
TE:  Hacienda  Comelco  (LACM,  UCB). 

Lithurge  ( Lithurge ) chrysurus  (Fonscolombe) 

Litburgus  chrysurus  Fonscolombe,  1834:220.  2 <3. 

RANGE 

Countries  bordering  western  Mediterranean  Sea;  adventive 
in  eastern  United  States. 

BIONOMICS 

According  to  Roberts  (1978),  females  construct  communal 
nests  in  both  soft  and  hard  woods,  causing  serious  damage. 


10  Contributions  in  Science,  Number  343 


Snelling:  North  American  Lithurge 


Cells  are  usually  separated  by  wood-dust  partitions  and  are 
packed  with  a pollen  and  nectar  mass  gathered  from  Cen- 
taurea  maculosa  (Compositae),  an  introduced  weedy  plant. 
When  partitions  are  absent,  two  or  three  regularly  spaced 
larvae  feed  on  the  single  large  pollen  mass.  When  larval 
feeding  is  completed,  the  mature  larva  defecates  and  spins  a 
pupal  cocoon.  Immatures  overwinter  as  prepupae,  and  pu- 
pation occurs  during  the  following  spring. 

DISCUSSION 

Friese  (1911)  gives  a complete  synonymy  for  L.  chrysurus. 
Roberts  (1978)  reported  this  Palaearctic  species  from  New 
Jersey,  where  it  had  been  established  for  at  least  three  years; 
nesting  biology,  behavior,  and  immature  morphology  were 
all  described  by  Roberts.  In  North  America,  this  bee  is  pres- 
ently known  only  in  New  Jersey,  where  it  is  oligolectic  on 
Centaurea  maculosa,  a plant  also  native  to  countries  bor- 
dering the  western  Mediterranean  Sea. 

ACKNOWLEDGMENTS 

For  the  loan  of  specimens  utilized  in  this  study,  I am  indebted 
to;  P.  H.  Arnaud,  Jr.  (CAS),  G.  R.  Else  (BMNH),  D.  K. 
Faulkner  (SDM),  M.  Favreau  (AMNH),  S.  Frommcr  (UCR), 
J.  E.  Gillaspy  (TAI),  T.  Griswold  and  F.  D.  Parker  (GRIS, 
UTSU),  F.  F.  Hasbrouck  (ARSU),  the  late  P.  D.  Hurd,  Jr. 
(USNM),  L.  S.  Kimsey  and  R.  O.  Schuster  (UCD),  U.  N. 
Lanham  (UCOL),  E.  G.  Linsley  (UCB),  S.  J.  Merrit  (TAMU), 
C.  D.  Michener  (UKAN),  B.  Ratcliffe  (UNEB),  M.  S.  Was- 
bauer  (CDFA),  and  F.  G.  Werner  (UNAR). 

I am  especially  grateful  to  G.  R.  Else  for  his  assistance  in 
locating,  and  making  available,  the  type  of  L.  apica/is  litto- 
ralis. 

LITERATURE  CITED 

Berthold,  A. A.  1827.  Latreille’s  natiirliche  Familien  des 
Theirreiche.  Weimar,  610  pp. 

Brach,  V.  1979.  Notes  on  the  biology  of  Lithwgus  gibbosus 
Smith  in  Florida  (Hymenoptera:  Megachilidae).  Bull.  So. 
Calif.  Acad.  Sci.  77:144-147. 

Cockerell,  T.D. A.  1898.  New  and  little-known  Hymenop- 
tera taken  by  Prof.  C.H.T.  Townsend  and  Mr.  C.M. 
Barber  in  New  Mexico  in  1898.  Ann.  Mag.  Nat.  Hist. 
(7)  2:448-457. 

. 1902.  The  genus  Lilhurgopsis.  Entomol.  News  13: 

182. 

. 1911.  Bees  in  the  collection  of  the  United  States 

National  Museum.  2.  Proc.  U.S.  Natl.  Mus.  40:24 1-264. 
. 1917.  In  W.P.  Cockerell,  Collecting  bees  in  south- 
ern Texas.  Jour.  N.Y.  Entomol.  Soc.  25:187-193. 

. 1937.  Bees  collected  by  Mr.  and  Mrs.  J.L.  Sperry 

and  Mr.  R.H.  Andrews  in  Arizona.  Bull.  So.  Calif.  Acad. 
Sci.  36:107-1 10. 


Cresson,  E.T.  1875.  Report  upon  the  collections  of  Hy- 
menoptera, In  Wheeler,  Rept.  Geogr.  Geol.  Explor.  Surv. 
West  of  100th  Meridian,  vol.  5,  pp.  707-728. 

Ducke,  A.  1 907.  Contribution  a la  connaissance  de  la  faune 
hymenopterologique  du  nord-est  du  Bresil.  Rev.  Ento- 
mol. 26:73-96. 

Fonscolombe,  B.  1834.  Notices  sur  les  genres  d’Hyme- 
nopteres  Lithwgus  et  Philoxera.  Ann.  Soc.  Entomol. 
France  3:219-224. 

Fox,  W.J.  1902.  Lilhurgopsis,  a new  genus  of  bees.  Ento- 
mol. News  13:137-140. 

Friese,  H.  1908.  Die  Apidae  (Blumenwespen)  von  Argen- 
tina nach  den  Reisenergebnissen  der  Herren  A.C.  Jen- 
sen-Haarup  und  P.  Jorgensen  in  den  Jahren  1 904-1907. 
Deutsch.  Entomol.  Zeits.  1908:1-94. 

. 1911.  Apidae  1.  Megachilinae.  Das  Tierreich  28: 

1-440. 

Hurd,  P.D.,  Jr.  1979.  In  K.V.  Krombein  et  al..  Catalog  of 
Hymenoptera  of  America  north  of  Mexico.  Washington, 
D.C.  2:1 199-2209. 

Latreille,  P.A.  1825.  Families  naturelles  du  regne  animal. 
Paris,  570  pp. 

Michener,  C.D.  1944.  Comparative  external  morphology, 
phylogeny  and  a classification  of  the  bees  (Hymenop- 
tera). Bull.  Amer.  Mus.  Nat.  Hist.  82:151-326. 

. 1965.  A classification  of  the  bees  of  the  Australian 

and  South  Pacific  regions.  Bull.  Amer.  Mus.  Nat.  Hist. 
130:1-362. 

Mitchell,  T.B.  1927.  New  megachilid  bees.  Psyche  34: 104- 
121. 

. 1938.  The  genus  Lithwgus  (Hymenoptera:  Mega- 
chilidae). Psyche  45:146-155. 

. 1962.  Bees  of  the  eastern  United  States.  N.  Car. 

Agric.  Exp.  Sta.  Tech.  Bull.  152:1-557. 

Moure,  J.S.  1949.  Las  especies  Chilenas  de  la  sub-famiiia 
Lithurginae  (Hym.-Apoidea).  Arq.  Mus.  Paranaense  7: 
265-286. 

Parker,  F.D.,  and  H.W.  Potter.  1973.  Biological  notes  on 
Lithurgus  apica/is  Cresson  (Hymenoptera:  Megachili- 
dae). Pan-Pacific  Entomol.  49:294-299. 

Parks,  H.B.  1 930.  Notes  on  Texas  bees.  Bull.  Brookl.  Ento- 
mol. Soc.  25:263-264. 

Roberts,  R.B.  1978.  The  nesting  biology,  behavior  and  im- 
mature stages  of  Litharge  chrysurus,  an  adventitious 
wood-bonng  bee  in  New  Jersey  (Hymenoptera:  Mega- 
gachilidae).  Jour.  Kans.  Entomol.  Soc.  51:735-745. 

Rozen,  J.G.,  Jr.  1973.  Immature  stages  of  lithurgine  bees 
with  descriptions  of  the  Megachilidae  and  Fideliidae 
based  on  mature  larvae  (Hymenoptera,  Apoidea).  Amer. 
Mus.  Novitates  2527,  14  pp. 

Smith,  F.  1853.  Catalogue  of  hymenopterous  insects  in  the 
collection  of  the  British  Museum.  Part  1,  197  pp.  British 
Museum,  London. 

Submitted  20  May  1982;  accepted  17  December  1982. 


Contributions  in  Science,  Number  343 


Snelling:  North  American  Lithurge  11 


SERIAL  PUBLICATIONS  OF  THE 
NATURAL  HISTORY  MUSEUM  OF  LOS  ANGELES  COUNTY 


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# Contributions  in  Science,  a miscellaneous  series  of  technical  papers  describing  orig- 
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m Science,, 

® Science  Series,  long  articles  on  natural  history  topics,  generally  written  for  the  layman. 

Copies  of  the  publications  in  these  series  are  sold  through  the  Museum  Book  Shop.  A catalog 

is  available  on  request, 

SCIENTIFIC  PUBLIC  ATIONS  COMMITTEE 


Craig  C.  Black,  Museum  Director 
Donald  Chaput 

Daniel  M.  Cohen,  Committee  Chairman 
John  M.  Harris 
Charles  L,  Hogue 

Robin  A.  Simpson,  Managing  Editor 
Gary  D.  Wallace 
Edward  C.  Wilson 


Primed  a?  Ail  Sea  Pre 

ii-  kit.i.’ m i 


IB 


, e:  . 


. lac.,  Lawrence,  Kansas 

MHi- 


as 


Li  ipft 
lllkg 


iMIF 


THE  TERTIARY  SPONGES  APHROCALLISTES 
AND  EURETE  FROM 

WESTERN  WASHINGTON  AND  OREGON 
J.  Keith  Rigby1  and  David  E.  Jenkins2 


ABSTRACT.  Extensive  collections  of  sponges  from  the  Eocene  to 
Miocene  of  southwestern  Washington  and  from  the  Oligocene  and 
Miocene  of  northwestern  Oregon  consist  mainly  of  Aphrocallistes 
polytretos  n.  sp.  This  new  sponge  has  irregular  undulating  to  tubular 
walls  that  look  like  liny-celled  honeycombs  because  of  their  coarse 
canals.  The  upper  cellular  part  of  the  skeleton  is  made  of  flattened 
hexiradiates,  but  the  basal  part  is  irregularly  dictyonine. 

The  less  common  Eurete goedertin.  sp.,  a branching  zigzag  tubular 
sponge,  occurs  in  the  Oligocene  part  of  the  Lincoln  Creek  Formation 
in  southwestern  Washington.  Root  tufts  of  large  monaxial  spicules 
occur  in  concretions  with  both  Eurete  and  Aphrocallistes  but  may 
not  be  part  of  either  of  these  sponges. 

Distributions  of  living  species  of  both  genera  suggest  by  analogy 
that  the  fossils  lived  at  a depth  of  300-350  meters.  Most  occur  in 
coarse  clastic  units  or  in  calcareous  concretions  with  a mixed  diverse 
fish-echinoid-sponge  assemblage.  This  fauna  may  have  been  swept 
into  still  deeper  water  by  pulses  of  high-energy  currents,  for  most 
sponges  and  echinoids  appear  fragmented.  Only  the  root  tufts  appear 
to  have  grown  in  place. 

INTRODUCTION 

Numerous  fossil  sponges  have  been  recovered  from  concre- 
tions eroded  out  of  relatively  deep-water  marine  Eocene  to 
Oligocene  sediments  near  Knappton  (Localities  2,  5-7,  Fig- 
ure 1),  from  Eocene  limestone  in  a quarry  near  Chinook 
(Locality  1,  Figure  1)  in  southwestern  Washington,  from 
Miocene  concretionary  beds  at  Astoria,  and  from  Oligocene 
deposits  near  Yaquina  (Locality  4,  Figure  1)  in  northwestern 
Oregon.  Near  Knappton,  the  major  locality,  the  sponges  are 
apparently  scattered  through  the  formation  but  are  most 
common  in  a zone  where  sponge-bearing  concretions  are 
particularly  abundant.  The  major  sponge  fauna  occurs  pre- 
sumably below  beds  that  have  produced  the  mollusks  being 
studied  by  Ellen  James  Moore,  of  the  U.S.  Geological  Survey, 
and  above  beds  that  have  produced  the  silicified  micro- 
mollusks  (bivalves  and  gastropods)  being  studied  by  Carole 
Hickman,  of  the  University  of  California  Museum  of  Pa- 
leontology. Fossil  barnacles  have  been  described  by  Victor 
Zullo  (1982)  from  localities  both  above  and  below  the  major 
sponge  zone. 


The  fossils  come,  in  large  part,  from  concretions  weath- 
ering out  of  a Holocene  landslide  block  in  the  Lincoln  Creek 
Formation  and  occur  as  float  along  the  Columbia  River  ter- 
race. The  slump  block  is  in  the  sea  cliff  at  the  head  of  the 
bay  between  Knappton  and  Grays  Point.  If,  as  Moore  (per- 
sonal communication,  December,  1981)  suggests,  “there  is 
a certain  amount  of  stratigraphy  preserved  in  the  float  from 
the  landslide  block,”  the  sponges  then  are  stratigraphically 
just  below  the  main  mollusk-bearing  beds.  “Because  the 
landslide  moved  generally  parallel  with  the  strike  of  the  beds, 
a proper  stratigraphic  sequence  seems  to  have  been  main- 
tained,” according  to  Moore  (personal  communication,  Au- 
gust, 1982). 

The  designation  Lincoln  Formation  was  initially  proposed 
by  Weaver  (1912:10-12)  for  Oligocene  beds  of  western 
Washington.  Weaver!  1937)  later  suggested  retaining  the  name 
Lincoln  Formation  for  the  entire  middle  Oligocene  of  Wash- 
ington. Weaver  and  others  ( 1 944:592)  noted  the  Lincoln  beds 
overlie  the  Keasey  Formation  and  unconformablv  underlie 
the  Miocene  Astoria  Formation.  The  Lincoln  Creek  For- 
mation in  the  Knappton  area  occurs  in  an  east-dipping  homo- 
cline,  west  of  outcrops  of  the  Astoria  Formation,  in  the 
Portuguese  Point-Grays  Point  area  west  of  Frankfurt.  Wash- 
ington (Wells,  1979). 

The  name  Lincoln  Creek  Formation  was  proposed  by  Beik- 
man  and  others  (1967)  to  replace  the  Lincoln  Formation, 
which  has  been  preempted  several  times.  In  the  type  area, 
the  formation  consists  of  700-3000  m (2000-9000  feet)  of 
clastic  rocks  of  predominantly  Oligocene  age.  Lincoln  Creek 
beds  unconformably  overlap  Eocene  and  older  units  near 
Oligocene  paleotopographic  high  areas  but  rest  conformably 
on  Eocene  beds  within  the  basins.  The  Lincoln  Creek  For- 
mation is  overlain  by  the  Astoria  (?)  Formation  in  the  type 
region.  Molluscan  and  foraminiferal  correlations  suggest  that 
the  Lincoln  Creek  Formation  ranges  from  the  Late  Eocene 


1.  Brigham  Young  University,  Provo,  Utah  84602. 

2.  Union  Oil  Company  of  California,  2323  Knoll  Drive,  Ventura, 
California  93003. 


Contributions  in  Science,  Number  344,  pp.  1-13 
Natural  History  Museum  of  Los  Angeles  County,  1983 


ISSN  0459-8113 


Figure  1.  Index  maps  to  sponge  localities  in  southwestern  Washington  and  northwestern  Oregon.  Arrows  indicate  collecting  localities.  A, 
Ona  Beach  locality  (4,  LACMIP  6148)  south  of  Newport  and  north  of  Seal  Rocks  on  the  coast,  Yaquina  15-minute  quadrangle,  Lincoln 
County,  Oregon.  B,  Bear  River  locality  where  calcified  sponges  and  mollusks  occur  in  “reefoidaP'  limestone  (1,  LACMIP  5802).  Chinook  IVi- 
minute  quadrangle.  Pacific  County.  Washington.  C,  Knappton  localities  in  the  Lincoln  Creek  Formation  where  it  is  exposed  in  the  sea  cliff, 
(8,  LACMIP  5787,  Oligocene;  2,  LACMIP  5842,  Upper  Oligocene;  5,  LACMIP  5843,  Lower  Oligocene;  6,  LACMIP  5844,  Eocene;  7,  LACMIP 
5852,  Oligocene)  on  the  north  shore  of  the  mouth  of  the  Columbia  River,  Knappton  71/2-minute  quadrangle.  Pacific  County,  Washington. 


2 Contributions  in  Science,  Number  344 


Rigby  and  Jenkins:  Tertiary  Sponges 


Foraminiferal 

Molluscan 

WASHINGTON 

OREGON 

\ 

Stages 

Stages 

Knappton 

Area 

Bear  River  Area 

Seal  Rocks  Area 

c 

<D 

O 

o 

Saucesian 

Ne  wportian 

Astoria  (?)  Formation 

Astoria  (?)  Formation 

Astoria  (?)  Formation 

Piilarian 

7 

? 

Nye  Mudstone 

Unit  4 

Juanian 

c 5842 

o 

Yaquina  Formation 

Oligocene 

ro  Unit  3 

5852 

o 

6148 

Zemorrian 

Matlockian 

ll 

(D 

Q) 

- Unit  2 

U 5843 

c 

o 

o 

c 

_l 

00 

in 

Lincoln  Creek  Formation 

Alsea  Formation 

o 

© £ 

Unit  1 

5844 

a.  o 

Refugian 

Galvinian 

Q.  o 
=>  LU 

? 

? 

Siltstone  of  Cliff  Point 
5802 

Figure  2.  Stratigraphic  distribution  of  localities  in  the  Knappton,  Bear  River,  and  Yaquina  (Seal  Rocks)  areas  showing  relationships  of  the 
informal  units  of  James  and  Gail  Goedert.  Natural  History  Museum  of  Los  Angeles  County  localities,  and  biostratigraphic  zones  of  the  Tertiary 
section  (modified  from  Zullo,  1982;  Wells,  1979;  Snavely  and  others,  1975;  Armentrout,  1981;  Rau,  1981). 


to  latest  Oligocene  (Rau,  1958,  1964,  1981;  Armentrout, 
1975.  1977,  1981). 

James  L.  Goedert  (Zullo,  1982:2-3)  has  subdivided  the 
Lincoln  Creek  beds  in  the  Knappton  outcrops  into  four  in- 
formal units  (Figure  2).  Unit  1 is  a barnacle-bearing  concre- 
tionary unit  associated  with  the  abundant  Ll-shaped  trace 
fossil,  Tisoa.  Unit  2 is  characterized  by  sponges,  small  aturiid 
nautiloids,  many  decapod  crustaceans,  and  marine  vertebrate 
remains  associated  with  tisoid  burrows.  Unit  3 is  character- 
ized by  a glass  sponge  fauna  and  Unit  4 by  an  abundance  of 
marine  vertebrate  and  invertebrate  fossils,  but  few  sponges. 
Zullo  ( 1 982:2-3,  fig.  2)  concluded  that  lower  Unit  1 is  Upper 
Eocene,  Unit  2 is  Lower  Oligocene,  and  LJnits  3 and  4 are 
Upper  Oligocene  (Figure  2). 

The  sponges  noted  here  from  Oregon  are  reportedly  from 
the  Astoria  Formation  at  the  type  locality  in  Astoria  and 
from  the  Yaquina  Formation  near  Seal  Rocks.  Moore  ( 1 963) 
described  outcrops  and  faunas  of  Astoria  beds  in  the  type 
area  and  southward  along  the  Oregon  coast  for  approximately 
70  miles.  One  specimen  of  Aphrocallistes  has  been  figured 
from  the  type  area  of  the  Astoria  Formation  by  Moore  ( 1 963, 
pi.  32,  fig.  16),  but  the  precise  locality  from  which  the  early 
collection  was  made  is  uncertain.  No  additional  sponges  have 
been  reported  from  other  exposures  along  the  sea  cliff  be- 
tween Astoria  and  Newport.  However,  a few  sponges  have 


been  recovered  from  the  Yaquina  Formation  south  of  New- 
port, in  the  Seal  Rocks  area  (Figure  1). 

Moore  (1963: 1 2)  described  Astoria  beds  in  the  coastal  area 
as  mainly  marine  silty  shale  and  fine-grained  sandstone,  with 
mterbedded  siltstone,  coarse-grained  sandstone,  conglom- 
erate, and  tuff.  She  reported  that  concretions  up  to  a foot 
and  a half  in  diameter  are  common  in  some  places.  Such 
concretions  have  produced  nearly  all  the  sponges  from  the 
Astoria,  Yaquina,  and  Lincoln  Creek  localities.  Some  con- 
cretions contain  a varied  molluscan  fauna,  fragments  of  car- 
bonized wood,  many  with  teredinid  borings,  marine  mam- 
mal bones,  fish  bones  and  scales,  a varied  foraminiferal  fauna, 
decapod  crustaceans,  and  barnacle  fragments.  Many  concre- 
tions are  coarse-grained  glauconitic  sandstone  that  shows 
evidence  of  vigorous  transport.  These  assemblages  may  have 
been  mixed,  that  is  may  have  been  derived  from  a variety 
of  depositional  sites,  by  downslope  movement. 

The  sponges,  in  general,  are  delicate  and  silicified  and  can 
be  freed  from  the  calcareous  matrix.  Some  concretions  were 
partially  etched  in  dilute  acetic  acid  and  screened  for  recovery 
of  all  microorganisms.  In  general,  the  concretions  are  highly 
argillaceous,  often  sandy  and  glauconitic.  For  complete  re- 
moval of  matrix,  the  partially  etched  calcareous  muds  were 
gently  brushed  and  washed  from  the  sponge  material.  Some 
fine  cleaning  of  specimens  was  done  with  a microsandblaster 


Contributions  in  Science,  Number  344 


Rigby  and  Jenkins:  Tertiary  Sponges  3 


air-abrasive  unit.  Some  delicate  etched  specimens  were  hard- 
ened in  weak  acetone-soluble  plastic  solutions. 

LOCALITIES 

The  fossil  sponges  described  here  were  all  collected  by  James 
L.  and  Gail  H.  Goedert,  except  for  the  sponge  reported  by 
Ellen  J.  Moore  (1963:89,  pi.  32,  fig.  16),  and  are  deposited 
in  the  Natural  History  Museum  of  Los  Angeles  County,  In- 
vertebrate Paleontology  Section  (abbreviated  LACMIP). 

LOCALITY  1.  Limestone  in  the  siltstone  of  Cliff  Point  of 
Wells  (1979)  in  a quarry  in  the  bluff  on  the  south  side  of 
Bear  River,  2.2  km  (1.4  miles)  northeast  of  Goulter  Ranch, 
on  the  section  line  between  Sections  20  and  21,  T.  10  N.,  R. 

1 0 W.,  on  the  Chinook  VA-minute  quadrangle.  Pacific  Coun- 
ty, Washington.  LACMIP  locality  5802,  Upper  Eocene. 

LOCALITY  2.  Lincoln  Creek  Formation  in  association 
with  other  invertebrate  fossils,  mammal  and  fish  bones,  and 
plant  fragments  in  calcareous,  argillaceous  concretions  that 
range  from  fine-grained  to  coarsely  clastic,  from  a landslide 
area  at  the  head  of  the  bay  between  Knappton  and  Grays 
Point  in  the  center  N 'A  N ‘A  of  Sec.  9,  T.  9 N.,  R.  9 W..  on 
the  Knappton  7‘A-minute  quadrangle  in  Pacific  County, 
Washington.  The  locality  is  approximately  1.6  km  (1  mile) 
northeast  of  Knappton.  LACMIP  Locality  5842,  Upper  Oli- 
gocene. 

LOCALITY  3.  Astoria  Formation  at  Astoria,  as  cited  by 
Moore  ( 1 963:90)  at  her  locality  la,  from  collections  of  James 
D.  Dana  discussed  by  Dali  (1909:140),  Miocene. 

LOCALITY  4.  Sponges  from  the  Yaquina  Formation,  in 
fossiliferous  concretions  from  rocks  exposed  at  Ona  Beach, 
0.4  km  (0.25  mile)  south  of  the  mouth  of  Beaver  Creek,  west 
of  U.S.  Highway  101,  2 km  (1.3  miles)  north  of  the  com- 
munity of  Seal  Rocks,  just  west  of  the  northwest  corner  of 
Sec.  1 9,  T.  12  S.,  R.  1 1 W.,  Lincoln  County,  Oregon,  Yaquina 
15-minute  quadrangle.  LACMIP  Locality  6148,  Miocene. 

LOCALITY  5.  Lower  Oligocene  part  of  the  Lincoln  Creek 
Formation,  from  concretions  weathering  out  of  the  sea  cliff 
in  the  northwestern  part  of  the  bay  between  Grays  Point  and 
Knappton.  approximately  305  m (1000  feet)  south  and  430 
m (1400  feet)  east  of  the  northwest  corner  of  Sec.  9,  T.  9 N., 
R.  9 W.,  on  the  Knappton  7‘A-minute  quadrangle,  in  Pacific 
County.  Washington.  The  locality  is  approximately  300  m 
( 1 000  feet)  west  of  Locality  2,  and  300  m ( 1 000  feet)  northeast 
of  Locality  7.  LACMIP  Locality  5843,  Lower  Oligocene. 

LOCALITY  6.  LIpper  Eocene  part  of  the  Lincoln  Creek 


Formation,  concretions  weathering  out  of  the  sea  cliff  in  the 
western  part  of  the  bay  between  Grays  Point  and  Knappton, 
approximately  0.8  km  (0.5  mile)  northeast  of  Knappton;  122 
m (400  feet)  east  and  520  m ( 1 700  feet)  south  of  the  northwest 
corner  of  Sec.  9,  T.  9 N..  R.  9 W.,  on  the  Knappton  7‘A- 
minute  quadrangle.  Pacific  County,  Washington.  LACMIP 
Locality  5844.  Upper  Eocene. 

LOCALITY  7.  Oligocene  part  of  the  Lincoln  Creek  For- 
mation, collected  from  the  “sponge  zone”  in  concretions 
weathering  out  of  the  sea  cliff,  approximately  1.3  km  (0.8 
mile)  northeast  of  Knappton,  305  m (1000  feet)  south  and 
580  m (1900  feet)  east  of  the  northwest  corner  of  Sec.  9,  T. 
9 N.,  R.  9 W..  Knappton  7‘A-minute  quadrangle.  Pacific 
County,  Washington.  This  is  the  most  productive  sponge 
locality  of  those  cited  here.  LACMIP  Locality  5852,  Oligo- 
cene. 

LOCALITY  8.  Lincoln  Creek  Formation  on  the  Columbia 
River,  approximately  122  m (400  feet)  east  of  the  boundary 
between  Sections  8 and  9,  T.  9 N.,  R.  9 W.,  approximately 
0.8  km  (0.5  mile)  northeast  of  Knappton.  Knappton  7‘A- 
minute  quadrangle.  Pacific  County,  Washington.  A general 
Knappton  locality  of  older  collections  before  units  were  es- 
tablished. LACMIP  Locality  5787,  Oligocene. 

SYSTEMATICS 

Class  Hexactinellida  Schmidt,  1870 
Order  Hexactinosa  Schrammen,  1903 
Suborder  Scopularia  F.  E.  Schulze,  1885 
Family  Aphrocallistidae  Gray,  1867 
Genus  Aphrocallistes  Gray,  1858 

Aphrocallistes  polytretos  n.  sp. 

Figures  3-10,  13-17,  21 

DIAGNOSIS.  Branching  tubular  to  cuplike  or  irregular 
undulating  leaf-  or  ear-shaped  sponges  with  upper  walls  per- 
forated by  honeycomblike,  subprismatic  to  polygonal  pri- 
mary canals  in  full  diarhysis  and  open  on  both  sides  of  the 
wall.  Basal  attachment  and  lower  part  of  walls  irregularly 
dictyonme  with  fewer  canals.  Each  canal  of  upper  wall  out- 
lined by  a complex  net  of  siliceous  skeletal  strands  produced 
by  hexiradiate  beams  whose  rays  have  been  distorted  to  be 
more  or  less  in  the  plane  of  the  porous  canal  walls.  No  sculp- 


Figures  3 through  10.  Aphrocallistes  polytretos  n.  sp.  (3)  Basal  view  of  two  specimens  showing  dense  irregular  attachment  areas  grading  up 
into  distal  radially  canalled  area.  Paratype,  LACMIP  6601,  Locality  2,  XI.  (4)  Natural  vertical  section  through  an  irregularly  dictyonine  base 
and  associated  lower  canalled  area.  Definition  of  canals  increases  away  from  base,  along  with  increase  in  modification  of  spicule  structure 
toward  hexiradiates.  Paratype,  LACMIP  6602,  Locality  8,  X2.  (5)  Side  view  of  well-preserved  paratype  with  honeycomblike  wall  and  tubular 
to  cuplike  form  but  with  broken  upper  end.  Base  of  this  same  specimen  is  shown  in  upper  right  of  Figure  3.  LACMIP  6601,  Locality  2,  X2. 
(6)  Fragments  of  sponge  showing  coarse  conglomeratic  and  glauconitic  nature  of  common  matrix.  LACMIP  6603,  Locality  7,  XL  (7)  Side 
view  of  broad  flattened,  moderately  fine-textured  paratype.  LACMIP  6604,  Locality  7,  XL  (8)  Side  view  of  holotype  showing  annular  tubular 
form  and  honeycombed  walls.  Reverse  side  has  concave-up.  dictyonine-spiculed  diaphragmhke  interruptions  that  probably  marked  pulses  in 
growth.  LACMIP  6600,  Locality  2,  XL  (9,  10)  Stereoscopic  photomicrographs  of  skeletal  net  of  the  holotype  showing  canal  patterns,  shape, 
and  hexiradiate  skeleton  typical  of  the  middle  and  distal  part  of  the  sponge.  LACMIP  6600.  Locality  2,  X5. 


4 Contributions  in  Science,  Number  344 


Rigby  and  Jenkins:  Tertiary  Sponges 


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Contributions  in  Science,  Number  344 


Rigby  and  Jenkins:  Tertiary  Sponges  5 


ture  is  evident,  except  for  the  undulating  surfaces.  Polygonal 
canals  are  generally  1.0- 1.5  mm  across  in  a wall  2-5  mm 
thick.  Upper  part  of  skeleton  of  irregular  flattened  hexiradiate 
dictyonal  beams  with  largest,  earliest  formed  beams  essen- 
tially horizontal  but  connected  with  irregularly  diagonal  and 
vertical  strands  in  a cross-bracing  crudely  hexiradiate  pat- 
tern, like  an  expanded  metal  screen,  with  roughly  hexagonal 
openings  often  subdivided  into  triangular  sections.  All  beams 
fused  into  a solid  network;  each  is  minutely  spinose. 

Canals  much  larger  and  less  oblique,  and  skeleton  in  ma- 
ture parts  much  more  hexiradiate-dominated  and  much  less 
clearly  dictyonine,  than  in  Aphrocallistes  vastus  whiteavesi- 
anus  Lambe  and  A.  vastus  vastus  Schulze.  Canals  much 
larger  but  more  closely  spaced  and  more  commonly  prismatic 
than  in  A.  alveolites  Roemerand  A.  cylindrodactylus  Schram- 
men. 

DESCRIPTION.  Numerous  nearly  complete  siliceous 
fragments  to  broken  bits  of  the  sponge  occur,  often  as  several 
pieces  with  different  growth  forms  in  single  concretions.  The 
holotype  is  a subtubular  undulating  sponge  with  a diameter 
of  30-35  mm.  It  is  approximately  80  mm  tall  and  is  sub- 
divided into  three  subequal  sections  with  saucer-shaped, 
noncanalled  partitions  with  a texture  like  that  of  bases  on 
other  specimens. 

The  wall  expands  upward  from  a thickness  of  2. 0-2. 5 mm 
in  the  lower  part  of  the  fragment,  to  2. 5-3. 5 mm  in  the  middle 
segment,  and  to  a maximum  of  4. 0-4. 5 mm  at  the  upper  end 
of  the  fragment.  The  arcuate  saucer-shaped  noncellular  par- 
titions are  generally  0. 5-1.0  mm  thick  in  the  center  of  the 
tubular  opening.  The  lower  segment  is  approximately  1 5 mm 
high,  the  middle  segment  is  20  mm  high,  and  the  uppermost 
is  20  mm  high,  above  the  uppermost  cellular  separation. 
Other  fragments  show  the  same  range  in  wall  thickness,  al- 
though most  are  only  undulating  irregular  platelike  bits  or 
tube  sections.  Some  nearly  complete  cup-shaped  sponges  show 
the  massive  base,  a few  millimeters  across,  and  a tubular 
form  8-30  mm  in  diameter  and  up  to  90  mm  long  or  high. 

Individual  canals  pierce  the  walls,  essentially  normally,  in 
full  diarhysis.  They  are  1.0-1. 8 mm  across  on  the  outside 
but  are  only  0.8-1. 5 mm  across  on  the  inside,  showing  ex- 
pansion of  the  canals  related  to  their  radial  orientation.  They 
are  closely  packed,  although  not  in  predictable  horizontal  or 
vertical  series,  and  are  so  spaced  that  7-8  occur  in  10  mm. 
both  horizontally  and  vertically  on  the  exterior.  Many  of  the 
canals  have  rectangular  cross  sections,  but  with  rounded  mar- 


gins. Hexagonal  and  pentagonal  openings  also  occur,  com- 
monly in  the  middle  unmodified  parts  of  the  segments.  All 
canals  have  porous  walls  so  that  there  could  have  been  com- 
plete interconnection  with  adjacent  ones.  Canal  walls  are 
perforated  at  irregular  intervals  with  distinct  rounded  open- 
ings 0.10-0.22  mm  across.  These  are  less  common  than  the 
much  smaller  and  more  irregularly  sized  triangular  openings 
that  occur  between  the  diagonal  and  vertical  beams  in  the 
hexiradiate-based  skeletal  net. 

The  skeleton  is  composed  of  fused  solid  beams  that  are 
difficult  to  separate  into  single  spicular  elements.  Walls  are 
principally  outlined  by  horizontal  fibers  that  tend  to  bifurcate 
horizontally  at  the  wall  intersections  and  vertically  in  the 
middle  part  of  the  wall.  These  are  the  first  beams  secreted 
at  any  level  in  the  skeleton  and  are  the  most  massive  units. 
They  are  spaced  5-7  horizontal  beams  per  mm.  measured 
vertically  along  the  canal  wall.  Most  of  these  strands  are  0.06- 
0.08  mm  across,  but  in  some  areas  where  the  skeletal  net  is 
dense,  they  have  been  thickened  to  0.10-0.12  mm  in  di- 
ameter. These  dominantly  horizontal  units  are  crossbraced 
by  diagonal  and  vertical  units  in  the  basic  flattened  hexira- 
diate skeleton.  Diagonal  beams  are  next  to  horizontal  ones 
in  size  and  are  generally  0.03-0.06  mm  across.  Vertical  beams 
are  apparently  the  last  added  and  are  the  smallest,  ranging 
from  tiny  hairlike  spines  0.01  mm  across  up  to  rods  0.04- 
0.05  mm  in  diameter,  fused  at  both  ends.  Diagonal  beams 
are  less  continuous  than  horizontal  ones,  and  vertical  ones 
are  the  least  continuous.  However,  vertical  beams  can  be 
traced  completely  along  the  canal  wall,  in  some  areas,  parallel 
to  the  trend  of  the  canals. 

Most  skeletal  elements  are  shared  between  two  adjacent 
canals  in  the  thin  wall.  Elsewhere,  particularly  in  proximal 
parts  of  the  sponge,  double  walls  occur,  with  each  canal  hav- 
ing its  own  subparallel  fused  series  of  elements.  These  strands 
bifurcate  to  produce  triangular  or  polygonal  radial  openings 
along  the  common  juncture  between  three  or  four  adjacent 
canals. 

Beams  are  apparently  solid  and  appear  to  have  been  built 
by  enlargement,  with  layered  secretions  over  initial  small 
needles  or  spines.  These  needles  are  common  in  young  parts 
of  the  skeleton,  pointing  upward  and  horizontally  from  ad- 
jacent thicker  strands.  Some  beams  are  so  enlarged  that  they 
combine  to  produce  a nearly  solid  fused  wall  in  the  older 
parts  of  the  net.  Exchange  between  canals  is  moderately  lim- 
ited in  these  areas,  but  in  the  outer,  more  open-textured  parts. 


Figures  11  through  17.  Root  tufts  and  Aphrocallistes  polytretos  n.  sp.  Figures  11,  12.  Root  tufts.  (11)  Large  concretion  with  aligned  oxeate 
root  tuft  spicules  in  fine-grained  calcareous  siltstone,  which  has  undergone  some  bioturbation  (lower  right).  LACMIP  6611,  Locality  7,  Xl. 
(12)  Large  monaxial  root  tuft  spicules  in  fine-grained  siltstone  showing  packing,  shape,  and  general  preservation  that  indicate  they  have  not 
been  moved  after  formation  of  the  tuft.  LACMIP  6612.  Locality  5,  X2.  Figures  13  through  17.  Aphrocallistes  polytretos  n.  sp.  (13)  Photo- 
micrograph showing  irregular  spicules  of  basal  attachment  and  dense,  noncanalled  pattern,  in  left  center  of  Figure  17.  LACMIP  6605,  Locality 
5,  X10.  (14)  Part  of  a concretion  containing  Aphrocallistes  with  irregular  branching  growth  form  in  variously  colored  fine-grained  matrix, 
which  produces  blotchy  appearance.  Paratype,  LACMIP  6606,  Locality  2,  XL  (15)  Photomicrograph  of  lower  part  of  sponge  showing  thick 
canal  walls  and  change  from  dictyonine-based  skeleton  in  lower  right  to  hexiradiate  skeleton  in  upper  left.  Side  of  same  specimen  in  Figure 
4.  Paratype.  LACMIP  6602.  Locality  8,  X10.  (16)  Photomicrograph  of  dictyonine  part  of  basal  skeleton  of  paratype  illustrated  in  Figure  4,  in 
left  center,  but  rotated  90  degrees.  Somewhat  swollen  spicule  nodes  and  irregular  fabric  are  characteristic.  LACMIP  6602,  Locality  2,  X10.  (17) 
Irregular  basal  region  and  part  of  honeycomblike  tubular  wall  of  paratype.  Base  shown  enlarged  in  Figure  13.  LACMIP  6605,  Locality  2,  X2. 


6 Contributions  in  Science,  Number  344 


Rigby  and  Jenkins:  Tertiary  Sponges 


Contributions  in  Science,  Number  344 


Rigby  and  Jenkins:  Tertiary  Sponges  7 


interchange  could  have  been  effected,  even  through  double 
walls  where  two  skeletal  layers  separate  adjacent  canals. 

The  entire  surface  of  each  spicular  strand,  particularly  the 
thickened  ones,  is  covered  by  minute  conical  spines  0.005- 
0.01  mm  high  and  across.  Some  tiny  elements  extend  as  fine 
needles  from  small  conical  bases.  These  cones  and  spines  are 
generally  0.01-0.02  mm  apart  and  appear  to  be  scattered 
irregularly  over  the  entire  surface  of  the  major  strands. 

DISCUSSION.  Generic  and  family  placement  of  these 
sponges  is  based  on  their  peculiar  flattened  hexactine  hexi- 
radiate  skeleton.  The  sponges  superficially  appear  like  lithis- 
tids,  composed  of  orchoclad  dendroclones.  The  “clads”  are 
fused  to  “clads"  ofother  spicules  to  produce  a flattened  roughly 
hexagonal  series  of  openings  with  one  spicule  axis  essentially 
normal  to  the  long  dimensions  or  axes  of  the  canals,  but 
individual  lithistid  spicules  cannot  be  identified. 

The  large  canals  might  be  considered  as  peculiar  porous 
calicles,  such  as  in  the  sclerosponge  Merlia  or  the  chaetaetids. 
However,  in  sclerosponges,  the  calicles  rise  from  an  imper- 
vious base  rather  than  forming  perforate  walls  around  a sub- 
cylindrical  or  platelike  porous  sponge. 

Schrammen  ( 19 12:358),  in  a summary  table  of  stratigraph- 
ic ranges  of  Cretaceous  fossils  of  northwestern  Germany, 
noted  that  Aphrocallistes  ranges  from  the  Late  Cretaceous 
into  Recent.  He  also  listed  the  genus  from  Tertiary  rocks  of 
Russia,  Japan,  and  Oran,  but  cited  no  references  for  those 
occurrences.  Schrammen  (1912:219-222)  differentiated  three 
species  of  the  genus  on  general  body  shape,  wall  thickness, 
presence  or  absence  of  terminal  sieve  plates,  or  whether  the 
forms  are  many  branched  or  relatively  simple  structures.  In 
general,  the  German  species  have  canals  considerably  more 
widely  separated  and  distinctly  smaller  than  our  specimens. 

Schrammen  (1912:185)  summarized  the  observed  depth 
ranges  of  living  species  of  Aphrocallistes.  In  general,  their 
range  is  from  approximately  100-1700  m.  Reid  (1968:549) 
cited  occurrences  of  A.  vastus  as  shallow  as  27.5  m from 
Puget  Sound.  He  also  listed  other  occurrences  off  Oregon  and 
Washington  in  water  97  and  108  m deep.  Living  species 
described  by  Schulze  (1887)  occur  essentially  in  the  North 
Pacific,  from  localities  in  the  Philippine  Islands,  Japan,  the 
Aleutian  Islands,  and  along  the  coast  of  Oregon  and  Wash- 
ington, California,  and  Vancouver  Island.  Thus,  it  is  not 
surprising  to  find  Aphrocallistes  in  Eocene  to  Miocene  beds 
in  Oregon  and  Washington. 

Aphrocallistes  was  collected  from  Astoria  beds  at  Astoria, 
Oregon,  on  the  south  bank  of  the  Columbia  River  (Dana, 


1849).  That  specimen  was  noted  by  Dali  (1909:140)  and 
figured  by  Moore  (1963,  pi.  32,  fig.  16)  as  an  unidentified 
ophiuroid  (?)  as  an  incidental  occurrence  to  her  extensive 
treatment  of  the  Miocene  mollusks  from  the  Astoria  For- 
mation of  Oregon. 

Canal  patterns  in  living  Aphrocallistes  vastus  whiteavesi- 
anus  Lambe,  illustrated  by  Reid  (1964,  text-fig.  47),  and 
Aphrocallistes  vastus  vastus  Schulze  show  markedly  oblique 
canals  and  clearly  defined  dictyonine-based  skeletons  in  a 
moderately  rectangular  arrangement.  Development  of  hex- 
iradiate  patterns  is  more  clearly  shown  in  Aphrocallistes  bea- 
trix  Gray,  as  figured  by  Reid  (1964,  text-fig.  49b).  Even  in 
that  species,  however,  there  is  a moderately  oblique  orien- 
tation to  some  canals  in  the  gastral  region,  and  wall  thick- 
nesses tend  to  be  somewhat  greater  than  in  the  Tertiary  species 
from  Washington  and  Oregon.  Canal  dimensions  in  each  of 
these  are  considerably  smaller  than  in  the  fossil  A.  polytretos. 
There  is  striking  similarity  of  skeletal  arrangements  in  A. 
beatrix  to  that  of  the  Tertiary  fossils.  The  walls  are  somewhat 
thicker  in  the  Recent  species,  and  the  canals  tend  to  be  more 
circular  than  distinctly  prismatic,  such  as  in  the  fossil  forms. 
Such  rounded  canals  are  the  general  pattern,  as  well,  in  the 
Cretaceous  sponges  described  by  Schrammen  (1912:21 9-222). 
A.  polytretos  is  similar  to  Aphrocallistes  beatrix  in  having  the 
dictyonal  strands  obscure,  in  contrast  to  Aphrocallistes  vastus 
varieties. 

Aphrocallistes  beatrix  Gray  (1858:1  14-1  1 5)  was  described 
from  the  Azores  in  the  Atlantic  Ocean.  Some  of  the  limited 
modern  distribution  is  unquestionably  related  to  sampling 
difficulty  and  spacing  of  samples  in  the  100-1000  m range 
in  the  modern  oceans. 

Associated  benthonic  foraminifersand  mollusks,  including 
the  nautiloid  Aturia,  suggest  water  depths  between  300  and 
600  m,  which  is  also  within  the  range  of  modern  Aphrocal- 
listes. The  assemblage  may  have  been  transported  from  mod- 
erately shallow  waters  of  300-400  m and  accumulated  ulti- 
mately in  depths  as  great  as  2000-4000  m. 

Specific  characteristics  are  the  size  of  individual  canals  and 
skeletal  elements  and  the  general  growth  form  of  the  sponges. 

Our  species  is  fairly  variable  and  ranges  at  least  from  Late 
Eocene  to  Miocene.  These  Tertiary  sponges  show  a general 
increase  in  canal  size,  a decrease  in  canal  wall  thickness,  and 
a tendency  for  a more  prismatic  canal  pattern,  when  com- 
pared to  the  Cretaceous  species  from  northwestern  Germany. 

Schrammen  (191 2:220)  noted  a terminal  or  oscular  sieve- 
like  diaphragm  on  branches  of  A.  alveolites  Roemer  and  A. 


Figures  18  through  22.  Eurete  goederti  n.  sp.  and  Aphrocallistes  polytretos  n.  sp.  Figures  18  through  20  and  22.  Holotype  of  Eurete  goederti 
n.  sp.  LACMIP  6609,  Locality  7.  (18)  Photomicrograph  of  gastral  part  of  dictyonine  skeletal  net  showing  marked  regularity  of  enlarged  strands 
and  somewhat  smaller  cross-connecting  beams.  Spicule  nodes  are  little  swollen.  Figure  23  is  a drawing  of  part  of  the  net  in  the  upper  left.  X5. 
(19)  Holotype  of  Eurete  goederti  and  associated  fragments  of  Aphrocallistes  in  bioclastic  matrix  of  a concretion  from  Locality  7.  The  regularly 
spaced  zigzag  pattern  of  the  branching  sponge  is  clearly  shown.  Proximal  is  to  upper  right  and  distal  to  lower  left,  where  circular  cross  section 
shows  near  shadowed  boundary  (arrow).  XL  (20)  Photomicrograph  of  thin  walls  of  the  holotype,  with  diverging  walls  that  are  3 or  4 spicule 
complexes  thick.  Gastral  surface  is  to  the  right.  X5.  (22)  Well-preserved  part  of  the  skeleton  showing  thickness  of  the  walls  in  lower  part  and 
regular  nature  of  skeleton  in  middle  and  upper  part.  Parts  of  three  dichotomous  branches,  producing  zigzag  structure  of  the  sponge,  are  shown. 
X3.  Figure  21.  Aphrocallistes  polytretos  n.  sp.,  paratype  illustrated  in  Figure  1 7,  here  enlarged  to  show  smaller  hexactine-based  spicules  partially 
filling  canals.  LACMIP  6605,  Locality  2,  XI  5. 


8 Contributions  in  Science,  Number  344 


Rigby  and  Jenkins:  Tertiary  Sponges 


Contributions  in  Science,  Number  344 


Rigby  and  Jenkins:  Tertiary  Sponges  9 


cylindrodactylus  Schrammen.  Diaphragms  are  present  in  the 
Washington  Oligocene  specimens,  but  in  general  these  are 
concave-up  divisions  within  tubular  parts  of  the  sponge  and 
must  mark  pulses  in  growth  of  the  sponge  rather  than  convex- 
up  terminations  like  those  on  the  German  Cretaceous  sponges. 
Orientation  in  the  Tertiary  sponges  is  well  defined  by  de- 
velopment of  irregular  basal  areas,  just  as  the  terminations 
are  well  defined  in  Schrammen’s  A.  cylindrodactylus  by  the 
rounded  fingerlike  growth. 

TYPE  SPECIMENS  AND  AVAILABLE  MATERIAL. 
Holotype  LACMIP  6600  from  Locality  7.  the  type  locality 
of  the  species,  and  paratypes  LACMIP  6601,  6603,  6607, 
6608  from  Locality  7,  LACMIP  6605,  6606  from  Locality 
5,  LACMIP  6604  from  Locality  7,  LACMIP  6602  from  Lo- 
cality 8.  In  addition  22  specimens  or  concretions  were  ob- 
tained from  Locality  1.  18  from  2,  1 from  4,  1 from  5,  2 
from  6,  and  12  from  Locality  7.  Some  concretions  contain 
several  specimens  or  fragments. 

ETYMOLOGY.  Poly,  Gr.,  many;  tretos,  perforated,  re- 
ferring to  the  many  coarse  canals  through  the  cellular-ap- 
pearing wall. 

Family  Euretidae  Zittel,  1877 
(fide  Schulze,  1887) 

Subfamily  Euretinae  Reid.  1958 
Genus  Eurete  Semper,  1868 

Eurete  goederti  n.  sp. 

Figures  18-20,  22,  23 

DIAGNOSIS.  Dendroid  or  irregular  zigzag  heteroto- 
mously  branching  axial  tube  with  short  lateral  branches,  both 
approximately  8-10  mm  in  diameter;  branches  approxi- 
mately 20  mm  apart.  Walls  0. 5-1.1  mm  thick;  skeletal  net 
euretoid,  lacking  prominent  ostia  and  postica;  moderately 
open  skeleton  lacking  well-defined  gastral  and  dermal  cortex; 
strands  originate  on  gastral  surface  and  parallel  that  surface 
before  arching  steeply  through  the  wall.  Gastral  strands  ap- 
proximately 0.10-0.23  mm  apart  and  0.10  mm  in  diameter, 
with  nodes  0.16-0.18  mm  in  diameter  spaced  0.4-0. 5 mm 
apart  along  strands  and  cross  connected  by  beams  0.06-0.09 
mm  across  to  form  rectangular  meshes.  Nodes  slightly  to 
distinctly  swollen  but  not  spherical. 

Branches  more  closely  and  regularly  spaced  and  slightly 
larger  than  in  E.  lithodendron  Reid,  and  much  better  devel- 
oped than  closely  spaced,  rudimentary  branches  in  E.  (?) 
setosum  Reid.  Lacks  cortex  and  ostia  that  are  developed  in 
both  E.  lithodendron  and  E.  (?)  setosum.  Growth  form  and 
branches  more  regular  than  in  E.  schmidti  treubi  Ijima. 
Spherical  nodes  absent  at  spicule  centra,  although  common 
in  related  species. 

DESCRIPTION.  Several  fragments  are  in  the  collection. 
The  most  complete,  the  holotype,  is  a fragment  approxi- 
mately 1 2 cm  high.  It  is  an  elongated  branched  form  in  which 
the  axial  tube  has  undergone  heterotomous  division  to  pro- 
duce a dichotomous  structure  in  which  one  of  the  two  branch- 
es ceases  to  grow  and  the  other  grows  to  divide  again.  This 


produces  a zigzag  growth  pattern  with  angles  of  approxi- 
mately 120-130  degrees  between  segments  of  the  axial  tube. 
The  axial  tubes  and  branches  are  8-10.5  mm  in  diameter. 
The  short  lateral  branches  are  spaced  approximately  20  mm 
apart  along  the  axial  tube  and  are  approximately  1 cm  long. 
Oscula  occur  at  the  ends  of  the  lateral  rudimentary  branches 
and  range  from  openings  to  somewhat  constricted. 

Walls  of  tubes  and  branches  are  0.5- 1.1  mm  thick  and 
have  a generally  thin,  but  well-defined  gastral  layer  of  thick- 
ened strands  and  beams.  Most  of  the  wall  is  composed  of 
relatively  open-textured  endosomal  skeleton.  A dermal  layer 
is  not  differentiated  in  the  well-developed  dictyonal  frame- 
work. The  thickened  gastral  layer  usually  affects  only  strands 
and  beams  at  the  immediate  gastral  surface. 

Neither  ostia  nor  postica  are  clearly  defined,  although  some 
irregular  interruptions  in  the  outer  part  of  the  skeleton  may 
represent  ostia.  These  are  subcircular  openings,  0. 1 5-0.3  mm 
across,  and  are  irregularly  developed  in  the  visible  exterior. 
No  similar  interruptions  are  evident  in  the  inner  part  of  the 
skeleton,  and  water  must  have  passed  through  the  wall  via 
the  open  skeletal  meshes. 

Strands  of  the  dictyonal  skeleton  originate  near  or  at  the 
gastral  surface  and  parallel  that  surface  before  swinging 
abruptly  derntally.  They  terminate  as  unattached  conical  to 
spinelike  rays  at  the  outer  margin  of  the  wall,  where  they  are 
often  nearly  at  right  angles  to  the  dermal  surface.  In  the  gastral 
layer,  strands  are  parallel  and  spaced  0.15-0.4  mm  apart. 
They  are  closest  immediately  distal  to  insertion  of  a new 
strand,  often  by  “branching,”  and  are  farthest  apart  imme- 
diately proximal  to  the  insertion.  Spicular  nodes  are  spaced 
by  beams  0.4-0. 5 mm  long  along  the  gastral  strands  and  are 
connected  laterally  by  beams  0.10-0.25  mm  long,  measured 
horizontally  on  the  gastral  surface.  Beams  of  gastral  strands 
are  0.08-0. 1 1 mm  in  diameter  midway  between  nodes,  with 
most  0.10  mm  across.  Lateral  gastral  beams  are  0.06-0.09 
mm  across,  with  most  in  the  upper  end  of  that  range.  Nodes 
are  0.10-0.22  mm  in  diameter  but  without  much  swelling 
other  than  slight  flaring  where  rays  meet.  Nodes  throughout 
the  skeleton  are  not  spherical  but  appear  almost  rectangular. 
Most  such  nodes  are  0. 1 6-0. 1 8 mm  across  in  the  endosomal 
part  of  the  wall. 

Endosomal  and  dermal  beams  are  slightly  smaller  than 
gastral  ones,  with  common  diameters  of  0.06-0. 10  mm  and 
lengths  of  0.2-0. 3 mm,  thus  spacing  nodes  three-dimen- 
sionally  that  distance  within  the  wall.  This  part  of  the  wall 
is  less  linearly  organized  than  the  gastral  part  and  is  2-4 
spicules  thick.  Free  beams  form  only  low  rounded  knobs  or 
cones  on  the  gastral  surface  but  pointed  spines  on  the  dermal 
surface. 

Skeletal  mesh  openings  are  generally  rectangular  in  the 
inner  part  of  the  skeleton  but  become  much  more  irregular 
and  commonly  triangular  in  the  middle  and  outer  part  of  the 

wall. 

Adnate  small  free  hexactines  occur  commonly  throughout 
the  skeleton.  They  are  generally  attached  to  a beam  by  one 
ray,  and  the  other  rays  of  the  tiny  spicules  are  free.  They 
range  from  small  spicules  with  rays  only  0. 1 5 mm  long  and 
0.005  mm  in  diameter  up  to  larger  forms  with  rays  0.3  mm 


10  Contributions  in  Science,  Number  344 


Rigby  and  Jenkins:  Tertiary  Sponges 


Figure  23.  Eurele  goederti  n.  sp.,  drawing  of  part  of  the  skeleton 
of  the  holotype  showing  smaller  attached  hexactines  and  larger  beams 
of  gastral  dictyonine  net.  Vertical  strands  are  dominant  elements  in 
this  part  of  the  net.  Distal  is  toward  the  top,  and  proximal  is  toward 
the  base.  LACMIP  6609,  Locality  7,  approximately  X40. 


long  and  0.008-0.0 10  mm  across  in  basal  ray  diameter.  They 
are  frequently  oriented  with  rays  parallel  to  the  major  dic- 
tyonal  elements,  in  the  general  plane  of  those  elements,  or 
at  approximately  45  degrees  to  those  elements  where  they 
project  out  into  the  “cubic”  meshes  (Figure  23).  Beams  in 
the  skeleton  all  show  granular  to  distinctly  spinose  surfaces, 
many  of  the  tiny  spines  only  0.005  mm  across  and  0.10  mm 
or  less  high. 

DISCUSSION.  Eurete  goederti  n.  sp.  is  most  similar  to 
Cretaceous  E.  lithodendron  Reid  (1961:34-38,  pi.  7,  figs.  4a, 
b)  in  general  growth  form  but  is  slightly  larger.  In  addition, 
branching  is  more  distant  in  E.  goederti  n.  sp.,  and  it  lacks 
a dermal  cortex  and  ostia,  which  are  present  in  E.  litho- 
dendron. Modern  E.  schmidti  treubi  Ijima  (1927:170;  Reid, 
1961:37,  fig.  2a)  also  has  a similar  growth  form  but  branches 
considerably  more  irregularly. 

E.  (?)  setosum  Reid  (1958:38-40),  from  the  Cretaceous  of 
Great  Britain,  is  also  a tubular  branching  form,  but  its  branches 
are  often  reduced  to  rudimentary  structures,  in  contrast  to 
the  distinct  branches  in  E.  goederti.  In  addition  E.  (?)  setosum 
has  a dermal  cortex  and  ostia  in  places,  and  branches  are 
spaced  relatively  closely  together.  Branching  in  E.  goederti 
is  considerably  more  regular  than  in  either  E.  lithodendron 
Reid.  E.  (?)  setosum  Reid,  or  in  the  living  E.  schmidti  treubi. 
The  northwestern  United  States  fossil  species  also  lacks 
spherical  spicule  nodes  that  are  common  in  the  other  species. 

Depth  ranges  of  living  species  of  Eurete  were  summarized 


by  Schrammen  (1912:1 84-185),  based  upon  work  by  Schulze 
and  Carter.  Shallowest  occurrence  cited  by  Schrammen  is 
220  m for  Eurete  carteri  Schulze,  and  deepest  is  717  m for 
E.  erectum  Schulze.  Most  species  are  listed  from  depths  of 
300  or  360  meters.  Reid  ( 1 968:549)  listed  Aphrocallistes  vas- 
tus Schulze  as  having  been  collected  in  depths  of  27.5  m 
(Puget  Sound),  97  m (Oregon),  and  108  m (Vancouver,  Brit- 
ish Columbia)  but  noted  that  the  Puget  Sound  site  is  the 
shallowest  authenticated  record  of  modern  dictyonine  hex- 
actinellids.  By  analogy  it  seems  likely  that  the  Washington 
Oligocene  species  had  a depth  range  of  100-350  m.  The 
fragmental  nature  of  our  material,  however,  suggests  that  the 
assemblage  may  have  been  transported.  Downslope  transport 
is  suggested  by  occurrence  of  these  and  associated  fossils  in 
thin,  coarse,  sandy  to  glauconitic  beds,  intercalated  as  high- 
energy  pulses  in  the  dominantly  fine-grained  sequence. 

TYPE  SPECIMENS  AND  AVAILABLE  MATERIAL. 
The  holotype,  LACMIP  6609,  is  the  largest  fragment  and 
occurs  with  Aphrocallistes  fragments,  pieces  of  wood,  fish 
fragments,  and  foraminifers  in  a coarse  sandy  siltstone  from 
Locality  7,  the  type  locality  of  the  sponge  species.  The  para- 
type  (LACMIP  6610)  and  an  additional,  more  fragmental 
piece  of  Eurete  are  also  from  the  same  locality.  One  concre- 
tion from  Locality  8 also  contains  tiny  scraps  of  dictyonal 
skeleton  that  are  probably  from  E.  goederti ; these  are  too 
small  to  determine  growth  habit  and  other  relationships  but 
have  the  same  proportions  in  the  gastral  net  as  the  holotype 
of  E.  goederti. 

ETYMOLOGY.  Goederti,  named  for  James  Goedert,  on 
whose  collections  this  and  parallel  studies  of  other  groups 
are  largely  based. 

Order,  Family,  Genus  Uncertain 

Hexactinellid  Root  Tufts 

Figures  11,12 

Several  concretions  contain  moderately  well  organized  root 
tufts  of  hexactinellid  sponges.  These  are  composed  of  con- 
centrically layered,  aligned,  large,  doubly  tapering  oxeas  (?), 
with  maximum  diameters  of  1. 1-1.4  mm  at  approximately 
midlength.  Neither  entire  length  nor  preserved  tips  were  ob- 
served on  any  single  spicule,  but  spicule  fragments  at  least  3 
or  4 cm  long  occur  in  fine-grained  calcareous  siltstone  where 
sharp  tips  and  double  taper  are  well  shown.  They  form  clus- 
ters 3 or  4 cm  across  and  are  commonly  isolated  from  other 
sponges.  In  one  small  concretion  from  Locality  2,  isolated 
large  root  tuft  spicules  occur  embedded  in  irregular  basal 
spicular  masses  of  Aphrocallistes.  Bases  of  other  specimens 
of  the  species  do  not  have  such  spicules  and  apparently  were 
cemented  to  bivalve  or  gastropod  fragments  or  pebbles.  This 
implies  that  the  tuft  spicules  are  not  an  integral  part  oi Aphro- 
callistes but  that  the  sponge  from  Locality  2 overgrew  ex- 
posed root  tuft  spicules,  which  formed  a solid  substrate 
“island”  on  the  muddy  bottom.  Concretions,  without  Aphro- 
callistes, from  the  same  locality  contain  both  root  tuft  clusters 
and  fragments  of  Eurete,  but  the  sponge  fragments  and  root 
spicules  are  not  mtergrown  and  may  be  unrelated  biologi- 
cally. The  root  tufts  may  represent  sponges  not  otherwise 


Contributions  in  Science,  Number  344 


Rigby  and  Jenkins:  Tertiary  Sponges  1 1 


preserved  in  the  collections.  Taxonomic  relationships  of  root 
tufts  are  unknown. 

FIGURED  SPECIMENS  AND  AVAILABLE  MATE- 
RIAL. Figured  specimen  LACMIP  6612  is  from  Locality  5, 
and  6611  is  from  Locality  7.  Five  other  concretions  con- 
taining tuft  fragments  were  collected  from  Locality  2,  and 
one  was  collected  from  Locality  5. 

Similar  root  tufts  of  large  bundled  oxeas,  which  show  con- 
centric layering,  occur  at  two  other  localities  (Armentrout. 
personal  communication,  1982).  One  such  tuft  was  collected 
by  Susan  Bee,  from  Portland  State  University,  from  Late 
Eocene  siltstone  of  the  upper  member  of  the  Keasey  For- 
mation. The  fossil  was  float  in  a 5-m  high  railroad  cut  south 
of  and  upslope  from  the  abandoned  railroad  trestle  across 
Highway  47,  between  Buxton  and  Vernonia,  approximately 
20  m (80  feet)  west  and  210  m (700  feet)  north  of  the  south- 
east corner  of  Sec.  8,  T.  3 N.,  R.  4 W.,  on  the  Vernonia  7 V2- 
minute  quadrangle,  Washington  County,  Oregon  (Locality 
Tok-Ml  12  of  Warren  and  others,  1945). 

The  other  tuft  was  collected  from  Late  Eocene  siltstone  of 
the  Lincoln  Creek  Formation  by  J.M.  Armentrout.  It  came 
from  the  upper  3-m  ( 10-foot)  interval  at  the  top  of  a cliff  on 
the  south  side  of  Canyon  River,  just  upstream  from  a logging 
road  bridge,  approximately  240  m (800  feet)  west  and  67  m 
(220  feet)  north  of  the  southeast  corner  of  Sec.  13,  T.  21  N., 
R.  6 W.,  on  the  Grisdale  15-minute  quadrangle.  Grays  Har- 
bor County,  Washington  (Locality  CR9  and  10  of  Armen- 
trout, 1973). 

ACKNOWLEDGMENTS 

The  sponges  and  associated  fossils  were  collected  by  James 
L.  and  Gail  H.  Goedert  over  a several-year  period.  Edward 
Wilson  of  the  Natural  History  Museum  of  Los  Angeles  Coun- 
ty suggested  the  project,  arranged  for  loans  of  the  collections, 
and  reviewed  the  manuscript.  Ellen  J.  Moore  of  the  U.S. 
Geological  Survey;  Lein  F.  Hintze  and  James  L.  Baer  of  the 
Department  of  Geology,  Brigham  Young  University;  Victor 
Zullo,  Department  ofGeology,  University  of  North  Carolina; 
and  John  M.  Armentrout  of  Mobil  Oil  Corporation,  Dallas, 
Texas,  critiqued  the  manuscript  and  added  stratigraphic  and 
biostratigraphic  information.  Armentrout  provided  data  on 
additional  occurrences  of  root  tufts  from  the  Keasey  and 
Lincoln  Creek  formations.  Drafts  of  the  manuscript  were 
typed  by  Ann  Bracken  and  Camille  Crezee.  David  Jenkins 
did  preliminary  work  on  part  of  the  collections  while  on  a 
graduate  internship  in  the  Department  of  Geology  at  Brig- 
ham Young  University.  Some  costs  of  manuscript  and  illus- 
tration preparation  were  covered  by  National  Science  Foun- 
dation grant  DEB  78-25229. 

LITERATURE  CITED 

Armentrout,  J.M.  1973.  Molluscan  paleontology  and  bio- 
stratigraphv  of  the  Lincoln  Creek  Formation,  late  Eocene- 
Oligocene,  southwestern  Washington.  (Ph.D.  disserta- 
tion) University  of  Washington.  Seattle,  478  pp. 

. 1975.  Molluscan  biostratigraphy  of  the  Lincoln 

Creek  Formation,  southwest  Washington.  In  Weaver, 


D.  (ed.).  Future  Energy  Horizons  of  the  Pacific  Coast; 
Paleogene  Symposium  and  selected  technical  papers. 
Annual  Meeting  of  the  Pacific  Sections,  American  As- 
sociation of  Petroleum  Geologists,  Society  of  Economic 
Paleontologists  and  Mineralogists,  Society  of  Economic 
Geologists,  Long  Beach,  California,  pp.  14-48. 

. 1977.  Cenozoic  molluscan  stages  of  Oregon  and 

Washington.  Abstracts  with  Programs.  Geological  So- 
ciety of  America  9(7):882— 883. 

. 1981.  Correlation  and  ages  of  Cenozoic  biostrati- 
graphic units  in  Oregon  and  Washington.  In  Armen- 
trout, J.M.  (ed.).  Pacific  Northwest  Cenozoic  Biostratig- 
raphy, Geological  Society  of  America  Special  Paper  1 84, 
pp.  137-148. 

Beikman,  H.N.,  W.W.  Rau,  and  H.C.  Wagner.  1967.  The 
Lincoln  Creek  Formation.  Grays  Harbor  Basin,  south- 
western Washington.  U.S.  Geological  Survey  Bulletin 
1244-1:11-11  14. 

Dali,  W.H.  1909.  Contributions  to  the  Tertiary  paleontol- 
ogy of  the  Pacific  coast,  I.  The  Miocene  of  Astoria  and 
Coos  Bay,  Oregon.  U.S.  Geological  Survey  Professional 
Paper  59,  278  pp. 

Dana,  J.D.  1849.  Geological  observations  of  Oregon  and 
northern  California,  U.S.  Exploration  Expedition  1 838— 
1842  under  the  command  of  Charles  Wilkes,  Geology, 
v.  10,  pp.  61  1-678;  Appendix,  pp.  722-723,  729-730; 
Atlas. 

Gray,  J.E.  1858.  On  Aphrocallistes,  a new  genus  of  Spon- 
giadae  from  Malacca.  Proceedings  of  the  Zoological  So- 
ciety of  London  26: 114-115. 

. 1867.  Notes  on  the  arrangement  of  sponges,  with 

a description  of  some  new  genera.  Proceedings  of  the 
Zoological  Society  of  London.  1867,  pp.  492-558. 

Ijima.  I.  1927.  The  Hexactmellida  ofthe  Siboga  Expedition. 
Siboga-Expeditie,  Leiden,  v.  6,  383  pp. 

Moore,  E.  J.  1963.  Miocene  marine  mollusks  from  the 
Astoria  Formation  of  Oregon.  U.S.  Geological  Survey 
Professional  Paper  419.  109  pp. 

Rau,  W.W.  1958.  Stratigraphy  and  foraminiferal  zonation 
in  some  of  the  Tertiary  rocks  of  southwestern  Washing- 
ton. U.S.  Geological  Survey  Oil  and  Gas  Investigations 
Chart  OC-57,  2 sheets. 

. 1964.  Forantinifera  from  the  northern  Olympic 

Peninsula,  Washington.  U.S.  Geological  Survey  Profes- 
sional Paper  375-G:G  1-G33. 

. 1981.  Pacific  Northwest  Tertiary  benthic  forami- 
niferal biostratigraphic  framework  — An  overview.  In 
Armentrout.  J.M.  (ed.).  Pacific  Northwest  Cenozoic  Bio- 
stratigraphy, Geological  Society  of  America  Special  Pa- 
per 184,  pp.  76-84. 

Reid,  R.E.H.  1958-64.  A monograph  of  the  Upper  Creta- 
ceous Hexactinellida  of  Great  Britain  and  Northern  Ire- 
land. Palaeontographical  Society,  London.  1958,  Part  I. 
pp.  i-xlvi;  1958,  Part  II.  pp.  xlvii-xlviii,  1-26;  1961, 
Part  III,  pp.  27-48;  1964,  Part  IV,  pp.  xlix-cliv. 

. 1968.  Bathymetric  distributions  of  Calcarea  and 

Hexactinellida  in  the  present  and  the  past.  Geological 
Magazine  1 05(  6):  546— 5 59 . 


12  Contributions  in  Science,  Number  344 


Rigby  and  Jenkins:  Tertiary  Sponges 


Schrammen,  A.  1912.  Die  Kieselspongien  der  oberen  Kreide 
von  Nordwestdeutschland;  II  Teil,  Triaxonia  (Hexacti- 
nellida).  Palaeontographica,  Supplement  5.  pp.  177-385. 

Schulze,  F.E.  1887.  Report  on  the  Hexactinellida  collected 
by  H.M.S.  Challenger  during  the  years  1873-76.  Report 
of  Scientific  Results  of  the  Voyage  of  the  H.M.S.  Chal- 
lenger. Zoology  21:1-513. 

Semper,  C.  1869.  Einige  neue  Kieselschwamme  der  Phil- 
ippen:  Hyalonema  Schultzei  n.  sp.  und  Eurete  n.  g. 
Verhandlungen  der  Physikalisch-Medicinischen  Gesell- 
schaft  in  Wurzburg,  new  series,  1 868,  1 : 29— 30  (also  pub- 
lished in  1 868,  Annals  and  Magazine  of  Natural  History 
2:372-373). 

Snavely,  P.D.,  Jr.,  N.S.  MacLeod.  W.W.  Rau,  W.O.  Addi- 
cott,  and  J.E.  Pearl.  1975.  Alsea  Formation  — An  Oli- 
gocene  marine  sedimentary  sequence  in  the  Oregon  Coast 
Range.  U.S.  Geological  Survey  Bulletin  1 395-F:Fl-F20. 

Warren,  W.C.,  R.M.  Grivetti,  and  H.  Norbisrath.  1945. 
Geology  of  northwestern  Oregon.  U.S.  Geological  Sur- 
vey Oil  and  Gas  Investigations.  Preliminary  Map  42. 


Weaver,  C.E.  1912.  A preliminary  report  on  the  Tertiary 
paleontology  of  western  Washington.  Washington  Geo- 
logical Survey  Bulletin  15:1-80. 

. 1937.  Tertiary  stratigraphy  of  western  Washington 

and  northwestern  Oregon.  Washington  University  (Se- 
attle) Publications  Geology  4:1-266. 

Weaver,  C.E. , and  others.  1944.  Correlation  of  the  marine 
Cenozoic  formations  of  western  North  America  (Chart 
no.  1 1 ).  Geological  Society  of  America  Bulletin  55:569- 
598. 

Wells,  R.E.  1979.  Geologic  map  of  the  Cape  Disappoint- 
ment-Naselle  River  area.  Pacific  County,  Washington. 
U.S.  Geological  Survey  Open-File  Report  79-389. 

Zullo.  V.A.  1982.  Arcoscalpellum  Hoek  and  Solidobalanus 
Hoek  (Cirripedia,  Thoracia)  from  the  Paleogene  of  Pa- 
cific County,  Washington,  with  a description  of  a new 
species  of  Arcoscalpellum.  Natural  History  Museum  of 
Los  Angeles  County,  Contributions  in  Science  336:1-9. 

Received  22  September  1982;  accepted  7 February  1983. 


Contributions  in  Science,  Number  344 


Rigby  and  Jenkins:  Tertiary  Sponges  13 


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Number  345 
7 October  1983 


* a mm  species  of 

AMERICAN  SALAMANDERS, /WITH, A REVIEW. 
DPICAL  GENERA 
IA,  CAUDATA,  PLETHGIKMIIJME) 


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eles.Gountsf.  * 900  Exposition  Boulevard  • Los  Angeles,  California  90007 


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Edward  C.  Wilson 


Primed  at  Allen  Press.  Inc.,  Lawrence.  Kansas 


NEW  GENERA  AND  A NEW  SPECIES  OF 
CENTRAL  AMERICAN  SALAMANDERS,  WITH  A REVIEW 
OF  THE  TROPICAL  GENERA 
(AMPHIBIA,  CAUDATA,  PLETHODONTIDAE) 


David  B.  Wake1 2 and 
Paul  Elias' 


ABSTRACT.  A new  genus  and  species  of  plethodontid  bolitoglos- 
sine  salamander  is  described  from  material  collected  in  northwestern 
Guatemala.  Bradvtriton  silus  new  genus,  new  species,  is  unique  in  a 
combination  of  structural  characteristics  that  includes  a laterally 
compressed  tail,  stocky  body  with  no  clearly  defined  neck,  and  short, 
slender  limbs  bearing  syndactylous  hands  and  feet.  To  diagnose  the 
new  genus,  an  analysis  of  the  entire  neotropical  assemblage  of  pleth- 
odontid  salamanders  was  undertaken.  Approximately  138  species 
belong  to  the  supergenus  Bolitoglossa  and  1 1 genera  are  recognized. 
The  genus  Chiropterotriton  is  shown  to  be  polyphyletic;  thus,  two 
new  genera  are  described.  Notolriton  new  genus,  includes  the  picadoi 
group  of  Chiropterotriton  beta.  Dendrotriton  new  genus,  includes  the 
bromeliacia  group  of  Chiropterotriton  beta.  Those  species  formerly 
called  Chiropterotriton  alpha  remain  as  the  sole  representatives  of 
the  genus.  Eight  of  the  eleven  neotropical  genera  are  shown  to  be 
monophyletic.  Of  the  three  exceptional  genera,  both  Dendrotriton 
and  Nototriton  are  nonparaphyletic  relative  to  all  genera  except  Oed- 
ipina,  but  Dendrotriton  is  easily  distinguished  from  Oedipina.  No- 
totriton may  be  paraphyletic  relative  to  Oedipina.  but  the  two  genera 
can  be  readily  distinguished  on  the  basis  of  major  differences  in 
ecology  and  shape.  Pseudoeurycea  comprises  morphologically  gen- 
eralized species  that  may  not  be  far  removed  from  the  morphology 
of  the  ancestral  stock  of  the  entire  neotropical  group.  Only  Nyctanolis 
and  Chiropterotriton  are  more  plesiomorphic  than  Pseudoeurycea. 
Lists  of  species  assigned  to  the  genera  are  provided.  Potential  rela- 
tionships within  the  supergenus  Bolitoglossa  are  discussed,  but  par- 
allelism and  convergence  have  been  so  extensive  that  no  definitive 
statement  concerning  generic  affinities  is  possible. 

RESUMEN.  Un  nuevo  genero  y especie  de  salamandra  plethodon- 
tida  bolitoglossina  se  describe  en  base  a material  recolectado  en  el 
noroeste  de  Guatemala.  Bradyiriton  silus.  nuevo  genero,  nueva  es- 
pecie, es  peculiar  por  su  combinacion  de  caracteres  que  incluye  una 
cola  comprimida  lateralmente,  un  cuerpo  macizo  con  cuello  esca- 
samente  definido,  y patas  delgadas  y cortas  provistas  de  manos  y 
pies  syndactilos.  Para  diagnosticar  este  nuevo  genero  se  realizo  un 
analisis  de!  grupo  completo  de  salamandras  plethodontidas  neotro- 
picales.  Las  aproximadamente  138  especies  pcrtenecen  al  superge- 
nero  Bolitoglossa,  en  el  cua!  se  reconocen  1 1 generos.  El  genero 


Chiropterotriton  se  sefiala  como  politiletico  y por  lo  tanto  dos  nuevos 
generos  se  describen.  Nototriton.  nuevo  genero,  incluye  el  grupo 
picadoi  Ac  Chiropterotriton  beta.  Dendrotriton,  nuevo  genero,  inclu- 
ye el  grupo  bromeliacia  de  Chiropterotriton  beta.  Las  especies  pre- 
viamente  incluidas  en  Chiropterotriton  alfa  permanecen  como  las 
unicas  represenlantes  de  este  genero.  Ocho  de  los  once  generos  neo- 
tropicales  se  senalan  como  monofileticos.  De  los  tres  generos  res- 
tantes,  tanto  Dendrotriton  como  Nototriton  son  monofileticos  en 
relacion  a todos  los  generos  excepto  Oedipina,  aunque  Dendrotriton 
es  facilmente  distinguible  de  Oedipina.  Nototriton  podria  ser  para- 
liletico  en  relacion  a Oedipina.  pero  estos  dos  generos  pueden  ser 
rapidamente  reconocidos  en  base  a sus  marcadas  diferencias  en  eco- 
logia  y forma  corporal.  Pseudoeurycea  comprende  especies  morfo- 
Iogicamente  generalizadas  que  podrian  estar  no  muy  alejadas  de  la 
morfologia  del  grupo  ancestral  del  ensamble  neotropical  analizado. 
Solo  Nyctanolis  y Chiropterotriton  son  mas  plesiomorficos  que  Pseu- 
doeurycea. Listas  de  las  especies  asignadas  a cada  genero  se  presen- 
tan.  Tambien  se  discuten  las  potenciales  relaciones  dentro  del  su- 
pergenero  Bolitoglossa.  pero  paralelismos  y convergences  han  sido 
tan  prevalentes  que  ninguna  opinion  definitiva  puede  plantearse  en 
cuanto  a alinidades  genericas. 

INTRODUCTION 

In  the  summer  of  1974,  the  junior  author  collected  several 
species  of  salamanders  in  a remote  area  in  northwestern  Gua- 
temala. Included  in  this  collection  were  three  species  that 
obviously  were  undescribed.  Subsequent  morphological 
analysis  showed  that  two  of  the  new  species  have  combi- 
nations of  traits  that  require  them  to  be  placed  into  two  new 
genera.  One  of  these  ( Nyctanolis ) has  been  described  else- 
where (Elias  and  Wake,  1983).  In  this  paper  we  describe  the 
second  new  genus.  We  also  report  the  results  of  a detailed 


1.  Museum  of  Vertebrate  Zoology  and  Department  of  Zoology, 
University  of  California,  Berkeley,  California  94720. 

2.  Research  Associate  in  Herpetology,  Natural  History  Museum 
of  Los  Angeles  County,  Los  Angeles,  California  90007. 


Contributions  in  Science,  Number  345,  pp.  1-19 
Natural  History  Museum  of  Los  Angeles  County,  1983 


ISSN  0459-8113 


analysis  of  all  of  the  currently  recognized  genera  of  tropical 
salamanders.  As  a result  of  this  analysis,  which  was  a nec- 
essary background  for  the  description  of  the  new  genera  and 
species,  two  additional  new  genera  are  erected.  These  are 
described  herein,  and  diagnostic  characters  are  given  for  all 
existing  genera  of  tropical  salamanders.  In  addition,  all  rec- 
ognized species  of  tropical  salamanders  are  referred  to  a ge- 
nus, and  relationships  among  the  genera  are  examined.  This 
analysis  reinforces  previous  views  (Wake,  1966;  Wake  and 
Lynch,  1976)  that  there  has  been  very  extensive  parallelism 
and  convergence  during  the  adaptive  radiation  of  pletho- 
dontid  salamanders  in  the  New  World  tropics. 

BACKGROUND 

Although  several  genera  of  New  World  tropical  salamander 
species  were  described  in  the  nineteenth  century,  and  some 
knowledge  of  the  diversity  of  the  group  was  available  at  that 
time,  the  most  authoritative  taxonomic  work  on  these  sal- 
amanders during  the  early  part  of  the  twentieth  century  (Dunn, 
1926)  placed  all  tropical  species  in  a single  plethodontid  ge- 
nus, Oedipus.  Dunn  considered  Oedipus  to  be  “a  large,  mod- 
em genus  of  some  30  species.  The  extremes  are  quite  different 
but  there  are  many  connecting  links.”  There  was  little  in- 
crease in  knowledge  of  tropical  salamanders  until  the  mid- 
I 930’s,  when  Schmidt,  Taylor,  and  other  workers  began  pub- 
lishing their  results  (for  historical  summary,  see  Wake,  1 972, 
and  Smith  and  Smith,  1976).  Taylor  (1940)  showed  that 
Oedipus  was  a preoccupied  name,  and  substituted  the  old 
name  Bolitoglossa  for  the  entire  assemblage,  except  for  a 
group  of  diminutive  species,  which  he  placed  in  Cope’s  ( 1 869) 
old  genus  Thorius.  Shortly  thereafter,  Taylor  (1944)  under- 
took a radical  revision  of  all  the  neotropical  salamanders,  in 
which  he  described  four  new  genera  (Chiropterotriton.  Par- 
vimolge,  Magnadigita,  Pseudoeurycea),  resurrected  Oedi- 
pina  and  Haptoglossa.  and  continued  to  recognize  Bolito- 
glossa and  Thorius.  Since  that  time,  there  has  been  relative 
stability  in  the  generic  classification  of  the  group.  Lineatriton 
was  established  by  Tanner  (1950),  Magnadigita  was  placed 
in  the  synonymy  of  Bolitoglossa  by  Wake  and  Brame  ( 1 963), 
and  Haptoglossa  was  placed  in  the  synonymy  of  Oedipina 
by  Brame  (1968).  The  entire  assemblage  was  characterized 
and  the  genera  defined  by  Wake  ( 1966),  who  established  the 
supergenus  Bolitoglossa  for  this  group.  This  supergenus,  and 
the  supergenera  Hydromantes  and  Batrachoseps,  were  in- 
cluded in  the  tribe  Bolitoglossini,  subfamily  Plethodontinae, 
of  the  family  Plethodontidae.  Many  species  have  been  dis- 
covered since  1926;  herein,  we  recognize  138. 

The  most  recent  comprehensive  treatment  of  the  super- 
genus Bolitoglossa! Wake  and  Lynch,  1976) dealt  mainly  with 
ecology,  distribution,  and  biogeography  and  offered  little  new 
morphological  or  taxonomic  analysis.  Since  the  publication 
of  that  paper,  major  new  discoveries  have  been  made;  here, 
we  attempt  to  give  phylogenetic  perspective  to  the  available 
information. 

The  present  effort  is  not  definitive,  for  continued  new  dis- 
coveries indicate  that  our  knowledge  of  the  group,  even  at 
the  generic  level,  remains  incomplete.  We  are  aware  of  many 


undescribed  species,  and  a number  of  described  species- 
including  some  crucially  important  ones— are  known  from 
only  one  or  a very  few  individuals.  Nevertheless,  we  believe 
that  the  broad  outlines  of  relationship  are  sufficiently  clear 
to  justify  the  substantial  revision  we  undertake  here. 

MATERIALS  AND  METHODS 

We  have  not  considered  in  detail  the  species  of  Bolitoglossa, 
Oedipina,  and  Thorius,  all  large,  monophyletic  genera,  which 
are  either  under  intense  study  presently  ( Bolitoglossa  by  D.B. 
Wake,  P.  Alberch,  A.  Larson,  and  colleagues,  Thorius  by  J. 
Hanken),  or  have  been  analyzed  recently  (Oedipina  by  Brame, 
1968).  Instead,  we  have  emphasized  apparently  polyphyletic 
and  paraphyletic  groups,  as  well  as  newly  discovered  species 
that  do  not  easily  fit  into  any  existing  genus.  In  these  critical 
cases,  we  have  analyzed  doubly  cleared  and  stained  speci- 
mens, histological  sections,  and  dissections  of  fixed  material. 
As  a point  of  departure  for  the  choice  and  analysis  of  char- 
acters, we  have  relied  on  the  literature,  which  will  be  cited 
where  appropriate. 

We  have  had  access  to  adequate  samples  of  most  of  the 
species.  Important  species  for  which  we  have  lacked  sufficient 
material  but  nevertheless  can  offer  useful  information  include 
(present  generic  designation  used):  Chiropterotriton  barbouri, 
C.  richardi,  and  Parvimolge  praecellens.  These  are  discussed 
in  appropriate  places  in  the  following  account. 

As  noted  above,  one  of  the  new  genera  has  recently  been 
described  (Elias  and  Wake,  1983).  We  begin  this  work  with 
an  account  of  the  characters  that  are  used  in  our  generic 
revision.  We  then  describe  a newly  discovered  genus  and 
species  and  erect  two  new  generic  names  to  encompass  pre- 
viously known  species.  Finally,  we  present  a preliminary 
phylogenetic  analysis. 

CHARACTERS  USED  FOR  ANALYSIS 

A necessary  first  step  in  the  process  of  phylogenetic  recon- 
struction is  the  identification  of  monophyletic  taxa  (sensu 
Eldredge  and  Cracraft,  1980).  Our  goal  is  to  deduce  mono- 
phyletic groups  from  a matrix  of  morphological  character 
states.  There  are  incomplete  data  for  too  many  species  to 
justify  an  extensive  analysis  at  the  species  level.  Accordingly, 
we  have  relied  on  the  literature  and  our  own  previous  work 
to  recognize  groups  of  species.  We  have  treated  these  groups 
as  hypotheses  and  have  investigated  whether  or  not  each  is 
a monophyletic  taxon.  A group  that  is  found  to  be  mono- 
phyletic is  then  treated  as  a genus.  We  believe  that  the  generic 
level  of  classification  should  combine  species  into  monophy- 
letic units  that  are  separated  from  other  such  units  by  mor- 
phological gaps.  Ideally,  the  gaps  will  coincide  with  ecological 
and  behavioral  differences  as  well.  Thus,  our  objective  is  to 
define  generic-level  units  that  are  meaningful  in  both  phy- 
logenetic and  ecological  terms. 

In  the  analysis  that  follows,  each  genus  is  scored  for  eigh- 
teen characters  (Table  1).  In  certain  genera,  there  is  some 
variation  in  these  characters  among,  or  even  within,  species. 
Only  if  a derived  condition  is  characteristic  of  every  known 
population  and  species  (i.e.,  only  if  it  appears  in  most  adults) 


2 Contributions  in  Science,  Number  345 


Wake  and  Elias:  Tropical  Salamanders 


is  the  genus  as  a whole  scored  as  derived.  Thus,  our  character 
scoring  should  reflect  the  most  primitive  common  denomi- 
nator for  each  genus  discussed  and  should  represent  the  sit- 
uation in  the  common  ancestor  of  the  included  species.  The 
disadvantage  of  this  conservative  approach  is  that  it  masks 
a certain  amount  of  parallelism. 

The  characters  used  in  our  analysis  are  described  below. 
All  of  the  characters  are  treated  as  two-state  characters,  with 
primitive  (plesiomorphic)  and  derived  (apomorphic)  states 
identified  on  the  basis  of  outgroup  analysis  (Eldredge  and 
Cracraft,  1980).  Our  outgroups  are  the  genera  Hydromantes 
and  Batrachoseps,  the  other  members  of  the  tribe  Bolito- 
glossini;  and  the  members  of  the  tribe  Plethodontini.  Most 
of  the  osteological  characters  have  already  been  discussed  in 
detail  by  Wake  (1966)  and  Lynch  and  Wake  (1978).  By  con- 
vention, we  code  the  characters  (0)  to  indicate  primitive  and 
(1)  to  indicate  derived  states. 

1.  Mesopodial  mineralization.  Mesopodials  are  cartilagi- 
nous throughout  life  (0)  or  are  mineralized  in  adults  (1). 

2.  Tibial  spur.  A well-developed  rodlike  structure,  free 
from  the  shank  of  the  bone,  is  present  proximally  (0)  or  is 
absent  or  reduced  to  a low  ridge  ( 1 ). 

3.  Carpal  fusion.  The  ulnare  and  intermedium  are  discrete 
(0)  or  fused  ( 1 ). 

4.  Tarsal  fusion.  Distal  tarsals  four  and  five  are  discrete 
(0)  or  fused  (1). 

5.  Mental  glands.  Clusters  of  individual  glands  form  a 
small  patch  located  near  the  tip  of  the  chin  in  male  tropical 
salamanders  (Truffelli,  1954).  The  patch  is  either  externally 
visible,  and  ovoid,  circular,  or  subtriangular,  with  rounded 
or  low  columnar  individual  internal  glands  (0),  or  is  exter- 
nally obscure,  with  long,  tubular  individual  internal  glands 
that  are  oriented  posteriorly  from  their  openings  near  the 
chin  (1).  The  tubular  glands  extend  posteriorly  as  twisted 
tubes  covered  by  the  skin  of  the  gular  area. 

6.  Stylus  of  operculum.  The  stylus,  or  columella,  is  not  well 
developed  in  any  of  the  tropical  species,  but  it  may  be  present, 
with  a distinct  rodlike  shape  (0)  or  reduced  to  a broad  bulge 
or  be  entirely  absent  (1). 

7.  Preorbital  processes  of  vomers.  These  slender  processes, 
which  may  or  may  not  bear  teeth,  are  either  present  between 
the  internal  nares  and  the  orbit  (0)  or  absent  (1). 

8.  Prefrontal  bones.  These  bones  are  either  present  in  vary- 
ing degrees  of  development  (0)  or  absent  (1). 

9.  Septomaxillary  bones.  These  bones  are  not  well  devel- 
oped in  any  tropical  salamanders,  but  small  ossicles  may  be 
present  (0)  or  absent  (1). 

10.  Frontal  processes  of  premaxillary  bone.  Where  only  a 
single  premaxillary  bone  is  present,  the  frontal  processes  are 
either  fused  together  at  the  point  of  origin  on  the  dorsal 
surface  of  the  pars  dentalis  and  for  some  distance  dorsopos- 
teriorly  (1)  or  separated  for  their  entire  length  (0).  We  score 
those  few  instances  in  which  the  processes  arise  separately 
and  subsequently  fuse  as  0. 

1 1 . Premaxillary  bones.  The  pars  dentalis  of  plethodontid 
salamanders  is  either  divided  (0)  or  fused  so  that  only  a single 
bone  is  present  ( 1 ). 

12.  Skull  roof.  The  parietal  bones  may  either  meet  or  ap- 


proach each  other  very  closely  on  the  midline,  thus  forming 
a complete  skull  roof  (0),  or  be  widely  separated  and  joined 
by  a connective  tissue  sheet,  thus  forming  an  incomplete  skull 
roof  ( 1 ). 

1 3.  Sublingual  fold.  A small  to  large  fold  of  glandular  skin 
underlying  the  tongue  may  be  present  (0)  or  absent  (1). 

14.  Limb  length.  Limbs  are  moderate  to  short  in  length, 
so  that  they  overlap  slightly  or  not  at  all  when  adpressed  (0), 
or  are  very  long,  so  that  they  overlap  by  more  than  two  costal 
interspaces  ( 1 ). 

1 5.  Tarsal  arrangement.  Distal  tarsal  four  is  discrete  from 
and  larger  than  distal  tarsal  five  and  articulates  with  the 
fibulare  (0),  or  distal  tarsal  five  is  discrete  from  and  larger 
than  four  and  articulates  with  the  centrale  (1).  When  distal 
tarsals  four  and  five  are  fused,  the  character  is  scored  as  0. 

16.  Trunk  vertebral  shape.  The  ratio  of  centrum  length  to 
posterior  centrum  diameter  is  low  (2.5-4),  and  the  vertebrae 
are  relatively  wide  and  short  (0),  or  the  ratio  is  high  (greater 
than  4),  and  the  vertebrae  are  relatively  narrow  and  elongate 
(1). 

17.  Number  of  trunk  vertebrae.  There  are  either  14  (0)  or 
18  or  more  (1). 

18.  Tail  shape.  The  tail  is  either  round  or  ovoid  in  cross 
section  (0)  or  strongly  compressed  laterally,  with  a dorsal, 
glandular  ridge  (1). 

The  eighteen  characters  are  listed  in  Table  1,  along  with 
the  states  that  characterize  each  of  the  1 1 genera  we  recognize 
in  this  paper.  This  table  alone  can  be  used  to  diagnose  all  of 
the  genera.  In  most  instances  the  genera  are  diagnosed  by 
autapomorphies  (derived  characters  unique  to  a given  taxon), 
combinations  of  synapomorphies  (derived  characters  shared 
by  two  or  more  taxa),  or  both.  In  some  instances,  determi- 
nation that  a given  genus  is  monophyletic  requires  an  analysis 
of  the  distribution  of  character  states  over  all  taxa;  such  cases 
are  detailed  in  the  Discussion.  All  1 1 genera  can  be  readily 
diagnosed  on  the  basis  of  unique  combinations  of  traits,  in- 
cluding both  apomorphic  and  plesiomorphic  states.  Bolito- 
glossa  probably  should  be  divided  into  at  least  two  taxa 
(Wake  and  Lynch,  1 976),  but  it  is  a well-defined  (if unwieldy), 
monophyletic  taxon  as  currently  recognized.  However,  our 
generic  concept  does  require  that  we  recognize  four  new  ge- 
neric taxa.  One  of  these,  Nyctanolis,  is  a recently  discovered 
monotypic  genus  that  has  been  described  elsewhere  (Elias 
and  Wake,  1983).  A second  newly  discovered  genus,  also 
monotypic,  is  described  immediately  below.  The  other  two 
new  genera  result  from  subdivision  of  an  existing  genus  and 
are  described  below  in  the  context  of  our  generic  diagnoses. 

DESCRIPTION  OF  A NEW  GENUS 
AND  SPECIES 

Bradytriton  new  genus 

TYPE  SPECIES.  Bradytriton  si/us  new  species. 

DIAGNOSIS.  A plethodontid  salamander  belonging  to 
the  subfamily  Plethodontinae,  tribe  Bolitoglossini,  superge- 
nus Bolitoglossa.  Bradytriton  is  a short-snouted,  stout-bod- 
ied, short-tailed  taxon  with  small  limbs  and  diminutive,  syn- 


Contributions  in  Science,  Number  345 


Wake  and  Elias:  Tropical  Salamanders  3 


Table  1.  Distribution  of  character  states  in  genera  of  neotropical  plethodontid  salamanders.  See  text  for  listing  of  characters.  0 = primitive 


state;  1 = derived  state. 


Genus 

Character 

1 

2 

3 

4 

5 

6 

7 

8 

9 

10 

1 1 

12 

13 

14 

15 

16 

17 

18 

Bolitoglossa 

0 

0 

0 

1 

0 

1 

0 

0 

1 

0 

1 

0 

1 

0 

0 

0 

0 

0 

Bradytriton 

0 

0 

1 

1 

1 

1 

0 

0 

1 

0 

1 

0 

0 

0 

0 

0 

0 

1 

Chiropterotriton 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

1 

0 

0 

0 

1 

0 

0 

0 

Dendrotriton 

0 

1 

0 

0 

0 

0 

0 

1 

0 

0 

1 

0 

0 

0 

0 

0 

0 

0 

Lineatriton 

1 

0 

0 

1 

0 

0 

0 

0 

1 

1 

1 

0 

0 

0 

0 

1 

0 

0 

Nototriton 

0 

0 

1 

1 

0 

1 

0 

0 

1 

1 

I 

0 

0 

0 

0 

0 

0 

0 

Nyctanolis 

0 

0 

0 

0 

0 

0 

0 

0 

1 

0 

0 

0 

0 

1 

0 

0 

0 

0 

Parvimolge 

1 

0 

] 

1 

1 

1 

1 

0 

1 

0 

1 

0 

0 

0 

0 

0 

0 

0 

Pseudoeurycea 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

1 

0 

0 

0 

0 

0 

0 

0 

Oedipina 

0 

1 

1 

1 

1 

1 

0 

1 

1 

1 

1 

0 

0 

0 

0 

0 

1 

0 

Thorius 

1 

0 

1 

1 

0 

1 

1 

0 

1 

0 

1 

1 

0 

0 

0 

0 

0 

0 

dactylous  hands  and  feet,  readily  distinguished  from  most 
other  members  of  the  supergenus  on  the  basis  of  those  fea- 
tures. It  differs  from  all  other  members  of  the  supergenus  in 
having  a laterally  compressed  tail  with  a relatively  massive 
dorsal  glandular  ridge.  Thorius  and  Parvimolge  have  short 
limbs,  but  these  genera  are  much  smaller  and  differ  in  other 
ways  as  well:  Thorius  has  an  incomplete  brain  case,  and  both 
genera  frequently  have  mineralized  mesopodial  and  hyo- 
branchial  ossifications  as  adults.  Nototriton  and  Dendrotriton 
have  long,  slender  tails.  All  Bolitoglossa  lack  sublingual  folds. 
Most  Pseudoeurycea  have  long  tails  and  long  limbs,  and  all 
lack  the  mesopodial  fusions  (ulnare-intermedium  in  manus; 
fourth  and  fifth  distal  tarsal  in  pes)  that  characterize  Bradytri- 
ton. 

ETYMOLOGY.  From  bradys,  Gr.,  referring  to  the  slow 
and  lethargic  movements  of  the  living  animals,  and  triton, 
Gr.,  a commonly  used  term  for  salamanders. 

Bradytriton  silus  new  species 

Figures  1 through  6 

HOLOTYPE.  Museum  of  Vertebrate  Zoology  (MVZ) 
131587,  an  adult  female  from  Finca  Chiblac,  15  km  NE 
Barillas,  Depto.  Fluehuetenango,  Guatemala,  elevation  1,310 
m (4,300  ft),  collected  by  P.  Elias,  6 September  1974. 

PARATYPES.  MVZ  131586,  LACM  134566,  same  data 
as  the  holotype;  MVZ  131589-131594  (6  specimens),  same 
locality  and  collector  as  holotype,  collected  7 September  1974; 
MVZ  1 34635-1 34637  (3  specimens),  El  Rayo,  elevation  1 ,370 
m (4,500  ft),  3 km  S buildings  of  Finca  Chiblac,  10  km  NE 
Barillas,  Huehuetenango,  Guatemala,  collected  by  J.  Jackson 
and  P.  Elias  on  1 September  1975;  MVZ  134638,  same  lo- 
cality and  collectors  as  preceding  series,  collected  on  31  Au- 
gust 1975;  MVZ  173063,  Finca  Chiblac,  10  km  NE  Barillas, 
Depto.  Huehuetenango,  Guatemala,  elevation  1,370  m (4,500 
ft),  collected  by  H.B.  Shaffer  and  P.  Elias  on  2 July  1977; 
MVZ  173064,  same  data  as  preceding  number,  collected  be- 
tween August  1975  and  October  1977  by  J.  Jackson  and  P. 
Elias. 


DIAGNOSIS  (measurements  in  millimeters).  See  Gener- 
ic Diagnosis.  A stout  species  of  moderate  size  (standard  length, 
SL,  in  four  adult  males,  39.1-53.3,  mean  45.3;  seven  adult 
females  44.5-53.0,  mean  49.0;  Table  2)  with  a very  stout, 
short  tail  (SL/tail  length  in  three  adult  males  is  1 .3-1 .7,  mean 
1.4;  in  six  adult  females,  1.2-1. 7,  mean  1.4),  short  limbs 
(when  adpressed,  fore  and  hind  limbs  leave  2-3,  mean  2.3, 
costal  interspaces  uncovered  in  four  adult  males;  2.5-3,  mean 
2.9,  in  seven  adult  females),  and  small,  syndactylous  hands 
and  feet.  The  relatively  broad  head  (SL/head  width  in  four 
adult  males  is  5.9-6. 1 , mean  6. 1 ; in  seven  adult  females,  5.7- 
6.6,  mean  6.2)  is  essentially  continuous  with  the  body,  and 
there  is  no  neck.  Maxillary  teeth  (41-50,  mean  44.0,  in  four 
adult  males;  40-59,  mean  47.9,  in  seven  adult  females)  and 
vomerine  teeth  (16-22,  mean  18.5,  in  four  adult  males;  17- 
25,  mean  20.6,  in  seven  adult  females)  are  moderate  in  num- 
ber. The  species  is  reddish  brown  in  color,  with  obscure 
streaking  and  flecking  of  lighter  and  darker  pigments. 

DESCRIPTION.  This  stout,  short-limbed  species  has  a 
very  short,  truncate  snout  and  diminutive  hands  and  feet. 


o I 2 3 4 


Figure  1.  Dorsal  view  of  MVZ  131587,  the  holotype  of  Bradytriton 
silus  new  genus  and  species. 


4 Contributions  in  Science,  Number  345 


Wake  and  Elias:  Tropical  Salamanders 


The  nostrils  are  small.  Labial  protuberances  are  poorly  de- 
veloped in  females,  which  have  an  especially  short  snout, 
but  are  large  and  wide  in  adult  males;  the  protuberances 
produce  a broad  muzzle  rather  than  the  elongate  snout  typical 
of  other  tropical  salamanders.  Mental  glands  are  present  in 


males  but  are  not  externally  demarcated;  the  glands  consist 
of  a cluster  of  moderately  long  tubules  that  converge  in  the 
region  of  the  mandibular  symphysis,  where  they  open  indi- 
vidually to  the  ventral  surface  of  the  anterior  mtermandibular 
region.  The  tongue  is  adetoglossal,  with  a distinct  small  pad. 


Figure  2.  Parasagittal  section  through  the  head  of  a specimen  (MVZ  134637)  of  Bradytriton  silus  new  genus  and  species.  The  section  is  near 
the  midline  and  passes  through  the  mandibular  symphysis. 

Left.  The  arrow  points  to  an  enlarged  premaxillary  tooth  and  indicates  the  part  of  the  section  that  is  magnified  in  the  view  on  the  right. 
When  the  mouth  is  closed,  the  premaxillary  teeth  extend  outside  the  mouth  and  lie  near  the  ventral  and  anterior  parts  of  the  lower  jaw,  near 
the  openings  of  the  mental  gland.  The  long,  tubular  ducts  of  the  mental  gland  are  evident  immediately  posterior  to  the  mandible.  Note  that 
no  external  ventral  swelling  is  present.  Also  well  displayed  in  this  section  is  the  complex  free  tongue  characteristic  of  the  tribe  Bolitoglossim. 
The  hyoglossal  muscles  are  attached  to  the  apparently  flexible  anterior  tip  of  the  basibranchial.  Immediately  anterior  to  the  tongue  is  the  large 
sublingual  fold,  characteristic  of  all  of  the  tropical  salamanders  except  Bo/itoglossa. 

Right.  Enlargement  of  the  premaxillary  tooth  at  the  end  of  the  arrow  in  the  left  part  of  this  figure.  The  pedicel  of  the  tooth  is  attached  to 
the  posteroventral  border  of  the  premaxillary  bone  (here  oriented  so  that  the  border  appears  to  be  ventral).  The  crown  is  relatively  large  and 
is  unicuspid.  The  cusp  is  strongly  hooked. 


Table  2.  Data  for  type  series  of  Bradytriton  silus  new  genus  and  species.* 


Sex 

Standard 

length 

Mead 

length 

Head 

width 

Foot 

width 

Hind 

limb 

length 

Fore 

limb 

length 

Tail 

length 

Maxil- 

lary 

tooth 

number 

Premax- 

illary 

tooth 

number 

Vomerine 

tooth 

number 

I.imb 

inter- 

val 

MVZ  131589 

6 

53.3 

11.3 

8.7 

3.3 

12.1 

9.4 

— 

50 

3 

17 

3.0 

MVZ  134636 

S 

46.4 

10.3 

7.6 

3.1 

1 1.0 

9.5 

36.0 

41 

5 

16 

2.0 

MVZ  134637 

6 

42.3 

oo 

bo 

6.8 

2.3 

9.6 

8.7 

31.6 

43 

6 

19 

2.0 

MVZ  134638 

6 

39.1 

9.0 

6.6 

2.3 

8.7 

8.0 

23.2 

42 

8 

22 

2.0 

MVZ  131591 

2 

53.0 

10.4 

8.7 

3.2 

10.7 

9.5 

30.6 

59 

8 

22 

3.0 

MVZ  131593 

2 

51.5 

10.3 

8.8 

2.5 

9.2 

9.6 

- 

48 

8 

20 

3.0 

MVZ  131586 

2 

50.1 

9.7 

7.8 

2.9 

1 1.0 

10.1 

40.4 

46 

9 

19 

3.0 

MVZ  134635 

2 

48.8 

10.1 

7.4 

2.7 

1 1.0 

9.2 

34.1 

46 

9 

25 

3.0 

MVZ  131587** 

2 

48.5 

9.5 

7.5 

2.5 

9.4 

9.5 

36.6 

52 

6 

17 

3.0 

MVZ  131592 

2 

46.6 

9.7 

8.2 

2.9 

1 1.0 

9.5 

34.5 

44 

10 

23 

2.5 

LACM  134566 

2 

44.5 

9.3 

6.8 

2.5 

8.6 

8.9 

33.3 

40 

9 

18 

3.0 

MVZ  131590 

J 

35.6 

8.1 

6.7 

2.2 

7.5 

7.3 

21.5 

31 

8 

16 

2.5 

MVZ  131594 

J 

34.4 

7.8 

6.1 

1.4 

7.2 

7.1 

22.3 

37 

8 

18 

2.0 

* All  measurements  are  in  millimeters. 
**  Holotype. 


Contributions  in  Science,  Number  345 


Wake  and  Elias:  Tropical  Salamanders  5 


10mm 


Figure  3.  Radiograph  of the  broken  tail  ofa  specimen  (MVZ  131586) 
of  Bradytriton  silus  new  genus  and  species.  Dorsal  to  the  top.  Note 
the  large  glandular  mass  above  the  vertebral  column. 

The  sublingual  fold  is  well  developed.  The  profile  of  the 
rounded  head  is  undifferentiated  from  the  trunk,  and  there 
is  no  apparent  neck  region.  A deep  unpigmented  groove  ex- 
tends beneath  the  eye,  following  its  curvature,  but  does  not 
extend  to  the  lip.  The  eyes  are  moderate  in  size  and  protrude 
slightly  beyond  the  margin  of  the  jaw.  The  postorbital  groove 
is  an  indistinct  furrow  that  extends  posteriorly  from  the  eye 
and  intersects  a deep  vertical  groove  at  right  angles.  The  latter 
groove  passes  behind  the  base  of  the  mandible  and  becomes 
a clearly  defined  nuchal  groove.  The  gular  fold  is  pronounced. 
Vomerine  teeth  are  few  to  moderate  in  number  and  are  ar- 
ranged in  a single,  curved  row  that  extends  laterally  beyond 
the  lateral  margin  of  the  internal  nares.  Maxillary  and  vo- 
merine teeth  increase  in  number  to  some  extent  with  in- 
creasing body  size.  The  maxillary  tooth  row  extends  back  to 


Figure  4.  Dorsal  views  of  the  left  foot  (left)  and  of  the  right  hand 
(right)  of  an  adult  male  (MVZ  1 73064)  Bradytriton  silus  new  genus 
and  species.  Cartilage  is  stippled.  The  phalangeal  elements  are  very 
poorly  ossified  and  are  less  distinct  than  illustrated  here  (see  text). 
Note  the  syndactylous  nature  of  the  digits. 


Figure  5.  Dorsal  view  of  the  skull  of  an  adult  male  (MVZ  1 73063) 
Bradytriton  silus  new  genus  and  species.  Bone  is  outlined,  and  car- 
tilage is  stippled.  The  external  nares  and  the  nasolacrimal  foramina 
are  black. 

a point  nearly  posterior  to  the  eyeball.  Premaxillary  teeth  are 
3-8,  mean  5.5,  in  four  adult  males,  6-10,  mean  8.4,  in  seven 
adult  females;  the  teeth  of  males  are  very  large  and  hooked 
and  protrude  under  or  through  the  lip  (Fig.  2).  The  trunk  is 
stout  and  cylindrical.  The  tail  is  of  moderate  length  and  is 
strongly  tapered  near  its  tip.  It  has  a marked  basal  constric- 
tion and  is  strongly  compressed  laterally.  The  tail  appears 
stout  when  viewed  laterally  but  seems  narrow  from  a dorsal 
perspective  (Figs.  I.  3).  Postiliac  glands  are  distinct.  The 
limbs  are  short  and  slender.  Hands  and  feet  are  diminutive 
and  syndactylous;  the  greatest  foot  width  is  about  ‘/15  SL  and 
the  foot  is  barely  wider  than  the  lower  limb.  The  digital  tips 
are  free,  but  adjacent  digits  are  fused  for  most  of  their  length 
(Fig.  4).  There  are  no  subdigital  pads.  The  toes,  in  order  of 
decreasing  length,  are  3,  2,  4,  5,  1;  the  fingers,  3,  2,  4,  1. 

MEASUREMENTS  OF  THE  HOLOTYPE  (in  milli- 
meters). Head  width  7.5;  snout  to  gular  fold  (head  length) 
9.2;  head  depth  at  posterior  angle  of  jaw  4.6;  eyelid  width 
1.9;  anterior  rim  of  orbit  to  snout  2.0;  horizontal  orbit  di- 
ameter 2.4;  interorbital  distance  2.3;  vomerine  teeth  virtually 
continuous  with  parasphenoid  tooth  patch;  snout  to  fore  limb 
12.6;  distance  separating  internal  nares  1.8;  distance  sepa- 
rating external  nares  1.9;  snout  projection  beyond  mandible 
0.3;  snout  to  posterior  angle  of  vent  (standard  length)  48.5; 


6 Contributions  in  Science,  Number  345 


Wake  and  Elias:  Tropical  Salamanders 


snout  to  anterior  angle  of  vent  44.4;  axilla  to  groin  28. 1 ; tail 
length  36.6;  tail  width  at  base  3.4;  tail  depth  at  base  4.8;  fore 
limb  length  (to  tip  of  longest  toe)  9.5;  hind  limb  length  9.4; 
hand  width  1.7;  foot  width  2.5. 

COLORATION  IN  LIFE.  The  dorsal  coloration  of  this 
species  is  reddish  brown.  This  color  is  broken  by  irregular 
Hecks  of  black  on  the  anterior  trunk  and  dominated  by  black 
on  the  head.  Brown  coloration  extends  halfway  down  the 
lateral  surfaces.  The  cheeks,  sides  of  the  tail,  and  lower  flanks 
are  black,  overlain  with  dense  white  flecks.  The  legs  are  brown 
proximally  but  grade  distally  to  black  with  white  flecking. 
The  feet  are  black  marked  with  white  flecks  dorsally.  The 
eye  is  chestnut  brown. 

COLORATION  IN  ALCOHOL.  The  dorsal  surfaces  of 
the  head  and  trunk  are  dark  gray.  This  color  gives  way  to 
lighter  gray  and  then  to  brown  on  the  tail.  Black  flecks  are 
superimposed  on  the  brown  at  the  tail  tip.  The  arms  and  legs 
are  dark  gray  with  some  light  flecks.  The  lips,  chin,  throat, 
and  undersides  of  the  legs  are  dark  gray  with  light  gray  flecks. 
The  ventral  surface  of  the  trunk  is  unmarked  dark  gray.  The 
ventral  surface  of  the  tail  lightens  distally  to  brown  inter- 
rupted by  darker  flecks.  The  palmar  surfaces  and  the  tips  of 
the  nasal  cirri  are  pale  gray. 

HABITAT.  This  species  lives  in  an  area  of  very  humid 
cloud  forest  that  received  5 to  6 meters  of  rainfall  annually. 
All  specimens  were  taken  within  2 km  of  the  type  locality 
and  were  found  in  direct  contact  with  the  substratum  under 
cover  objects.  Sympatric  with  Bradytriton  were  three  species 
of  Bolitoglossa,  one  species  of  Nyctano/is,  various  hylid  and 
leptodactylid  frogs,  lizards  of  the  genera  Anolis,  Sce/oporus, 
and  Lepidophyma,  and  one  member  of  the  snake  genus  Lep- 
todeira  (Elias,  in  press). 

OSTEOLOGY  AND  RELATED  MORPHOLOGY.  In- 
formation has  been  derived  from  two  cleared  and  stained 
male  specimens,  from  radiographs  of  the  entire  sample,  and 
from  histological  sections  of  the  head  and  neck  of  one  male. 

The  skull  (Fig.  5)  is  short  and  very  broad.  The  facial  portion 
of  the  skull  is  poorly  developed  and  is  little  expanded  in  front 
of  the  eyes.  The  anterior  cranial  elements  are  surprisingly 
weak  for  a moderate-sized  species,  and  the  bones  are  highly 
variable  in  shape,  position,  and  number  from  one  individual 
to  the  next,  and  even  from  one  side  to  the  other  in  the  same 
individual.  The  premaxillary  is  relatively  broad  and  has  a 
large  pars  dentalis.  However,  the  bone  virtually  “floats”  at 
the  anterior  end  of  the  skull;  in  most  individuals,  it  does  not 
contact  the  maxillaries,  and  in  some  it  is  separated  from  those 
bones  by  a sizeable  gap.  Even  when  a contact  exists,  the 
articulation  is  tenuous.  The  palatal  portions  of  the  premax- 
illary are  greatly  reduced  or  absent  and  do  not  contact  the 
anterior  extensions  of  the  vomers.  The  frontal  processes  arise 
separately  from  the  pars  dentalis  and  are  separated  for  their 
whole  length.  They  are  very  narrow  and  columnar  basally 
but  become  markedly  compressed  vertically  to  form  the  lat- 
eral margins  of  the  gland-filled  internasal  fontanelle.  The 
processes  are  generally  divergent  for  their  entire  length  and 
become  flattened  and  slightly  expanded  near  their  distal  tips. 
These  tips  overlap  the  expanded  facial  portions  of  the  frontals 
in  relatively  firm  (for  this  species)  articulations.  The  frontal 


Figure  6.  Dorsal  view  of  the  hyobranchial  apparatus  of  an  adult 
male  (MVZ  173063)  Bradytriton  silus  new  genus  and  species.  The 
entire  structure  is  cartilaginous.  The  paired  ceratohyals  ordinarily 
lie  closer  together  but  have  been  moved  somewhat  laterally  for  clarity 
of  illustration. 

processes  terminate  distinctly  posterior  to  the  posterior  tip 
of  the  nasals,  a little  behind  the  osseous  anterior  margin  of 
the  orbit.  The  mternasal  fontanelle  is  relatively  narrow,  ex- 
cept near  its  posterior  terminus.  In  three  adult  males  the 
nasals  are  strongly  protuberant,  extending  well  anterior  to 
the  jaw  outline,  but  they  are  only  slightly  to  moderately  pro- 
tuberant (Fig.  5)  in  the  other  males.  The  nasals  protrude 
slightly  in  two  females,  including  the  holotype.  Nasals  are 
very  irregular  in  size,  shape,  and  degree  of  ossification  but 


Contributions  in  Science,  Number  345 


Wake  and  Elias:  Tropical  Salamanders  7 


tend  to  be  roughly  triangular  to  quadrangular.  The  posterior 
tip  is  poorly  defined,  and  the  anterior  margin  is  very  irregular. 
Tiny  fragments  of  disconnected  bone  frequently  occur  around 
the  anterior  and  lateral  margins  of  the  nasal.  In  one  cleared 
and  stained  individual,  one  of  these  fragments  is  very  large 
and  is  situated  as  if  it  were  an  entirely  separate  bone  inter- 
calated between  the  nasal  and  the  prefrontal  (Fig.  5).  This 
fragment  is  only  slightly  smaller  than  the  prefrontal.  The 
nasals  only  slightly  overlap  the  anterior  parts  of  the  frontals. 
They  have  a variable  degree  of  contact  with  the  maxillary 
facial  processes  but  a very  narrow  contact  with  the  prefron- 
tals.  The  posterolateral  margin  of  the  nasals  may  be  slightly 
evacuated  for  the  passage  of  the  nasolacrimal  duct,  which 
passes  through  a shallow  but  distinct  channel  in  the  lateral 
part  of  the  prefrontal  and  enters  the  nasal  capsule  through  a 
foramen  in  the  anterior  margin  of  the  prefrontal.  The  fora- 
men may  form  a half-circle  in  the  prefrontal.  The  prefrontals 
are  about  one-third  the  size  of  the  nasals.  Some  small,  bony 
fragments  are  present  in  some  individuals  at  the  anteroven- 
tral  end  of  the  prefrontals.  The  prefrontals  are  relatively  long 
and  narrow  and  are  extensively  overlapped  by  the  facial  pro- 
cesses of  the  maxillary.  The  prefrontals  overlap  the  frontals 
only  slightly.  The  maxillaries  are  relatively  short  and  straight 
with  narrow,  pointed  anterior  and  posterior  projections  on 
the  dental  portion.  The  posterior  tips  extend  nearly  to  the 
posterior  margin  of  each  eyeball.  Maxillary  teeth  are  bicuspid 
and  moderate  in  size.  They  are  borne  along  all  but  the  tip  of 
the  dental  portion  of  the  maxilla.  The  relatively  well-devel- 
oped palatal  portion  of  the  maxillaries  is  separated  by  a strong 
articulation  from  the  lateral-most  part  of  the  vomerine  body. 
There  are  no  septomaxillary  bones. 

The  large,  morphologically  specialized  premaxillary  teeth 
of  males  are  attached  to  the  premaxillary  in  such  a way  that 
their  elongate  crowns  project  directly  anteriorly  and  pierce 
the  lip  (Fig.  2).  Once  through  the  lip,  the  unicuspid  (appar- 
ently only  the  lingual  cusp  is  retained)  makes  a sharp  90° 
turn  caudad  before  terminating  in  a point.  When  the  mouth 
is  closed,  the  tooth  tip  is  very  close  to  the  opening  of  the 
mental  gland.  This  gland,  which  has  a restricted  exit  at  the 
mandibular  symphysis,  is  a cluster  of  tubules  converging  to 
form  a small  cluster  of  apertures  (Fig.  2).  The  glands  and 
teeth  constitute  a functional  complex  that  serves  to  transfer 
the  glandular  secretion  to  females  during  courtship  in  a man- 
ner analogous  to  vaccination  (Arnold,  1977).  Teeth  of  similar 
shape  occur  in  some  species  of  Pseudoeurycea  (Taylor,  1941). 

The  large,  strong  vomers  are  in  limited  contact  only  at 
their  extreme  tips,  posterior  to  the  large  intervomerine  fon- 
tanelle.  A distinct,  narrow  process  at  the  anterior  end  of  the 
vomer  extends  toward  the  premaxillary  but  does  not  touch 
that  bone;  rather,  it  is  appressed  against  the  ventral  surface 
of  the  nasal  capsule  and  follows  that  structure  to  curve  up- 
wards at  its  anterior  tip.  Immediately  medial  and  anterior 
to  the  internal  naris,  the  body  of  the  vomer  is  strengthened 
by  a dorsal  bony  ridge  that  is  apparently  unique  among  boli- 
toglossine  salamanders.  The  large,  stout,  but  relatively  short, 
preorbital  process  falls  far  short  of  the  lateral  margin  of  the 
vomerine  body.  The  process  is  blunt  tipped,  and  it  has  a 
dorsal  dimension  that  is  very  unusual  in  that  it  seems  to 


become  continuous  with  the  antorbital  cartilage.  The  vo- 
merine teeth  are  borne  in  a single  curving  row  along  the  base 
of  the  vomer  body  and  nearly  to  the  tip  of  the  preorbital 
process.  The  bicuspid  teeth  are  about  the  size  of  the  maxillary 
teeth. 

Frontals  are  well  developed  and  articulate  firmly  with  each 
other  middorsally,  except  at  their  anterior  end.  The  posterior 
part  of  each  bone  is  a large,  broad  lobe  that  broadly  overlaps 
the  parietal.  A small,  anterior  lobe  of  the  parietal  abuts  the 
lateral  margins  of  the  posterior  lobe  of  the  frontal  but  does 
not  significantly  overlap  the  frontal.  The  facial  portion  of  the 
frontal  is  rather  poorly  developed.  Anteriorly,  the  frontal  is 
drawn  into  a narrow  point  lying  ventral  or  lateral  to  the 
frontal  processes  of  the  premaxillary. 

The  parietals  are  large  bones,  firmly  articulated  to  each 
other  and  to  all  surrounding  bones.  The  stout,  well-developed 
lateral  spurs  extend  anteroventrally,  overlapping  the  cartilag- 
inous insertion  of  the  ascending  process  of  the  palatoquadrate 
cartilage  into  the  braincase.  The  semicircular  canals  produce 
prominent  bulges  in  the  relatively  large  otic  capsules.  A small 
but  prominent  spinous  process  is  present  at  the  extreme  lat- 
eral margin  of  each  capsule.  A fibrous  mass  of  connective 
tissue  extends  from  this  process  to  the  quadrate.  The  squa- 
mosal lies  in  a distinct  depression  in  the  lateral  wall  of  the 
capsule.  The  large  parasphenoid  is  relatively  broad  and  blunt- 
tipped  anteriorly;  the  orbitosphenoids  are  relatively  widely 
separated.  Posterior  vomerine  teeth  are  in  two  bilateral  patches 
that  diverge  posteriorly.  The  patches  in  two  males  contain 
36-39  and  44-48  small,  bicuspid  teeth,  respectively.  The 
operculum  has  no  stilus,  although  a slight  protrusion  is  pres- 
ent. The  well-developed  quadrate  is  attached  to  the  otic  cap- 
sules by  relatively  large,  strong  squamosals.  A small  process 
extends  posteriorly  from  the  squamosal  and  is  connected 
by  a ligament  to  the  proximal  tip  of  the  ceratohyal. 

The  lower  jaw  is  rather  weak.  The  prearticular  is  relatively 
large,  but  low,  and  the  dentary  is  very  slender.  The  arc  of 
the  lower  jaw  is  relatively  flat  and  broad. 

The  hyobranchial  apparatus  is  typical  of  that  of  generalized 
members  of  the  supergenus  Bolitoglossa  in  being  cartilagi- 
nous and  lacking  a urohyal  (Fig.  6).  The  rather  slender  cer- 
atohyals  have  a discrete  and  well-developed  medial  process. 
The  very  narrow  anterior  process  is  drawn  into  a point  that 
extends  upward  into  the  sublingual  fold.  The  long  and  slender 
epibranchials  are  slightly  more  than  twice  the  length  of  the 
basibranchial.  The  basibranchial  is  slightly  less  than  twice 
the  length  of  the  second  ceratobranchial.  The  second  cera- 
tobranchials  are  much  stouter  than  the  slender  first  cerato- 
branchials.  The  first  ceratobranchials  are  relatively  well 
developed  in  comparison  with  those  of  other  tropical  pleth- 
odontids.  The  radii  of  the  first  basibranchial  are  continuous 
with  the  main  part  of  the  element,  and  there  is  no  sign  of  a 
fibrocartilage  joint.  The  broad-based  and  relatively  short  ra- 
dii are  joined  by  a distinct  connective  tissue  strand.  The  most 
distinctive  feature  of  the  entire  hyobranchial  apparatus  is  the 
very  well-developed  anterior  projection  that  is  unique  among 
the  tropical  genera  (Figs.  2,  6).  This  broad-based  and  rela- 
tively stout  projection  is  apparently  somewhat  flexible,  for 
the  cartilage  cells  at  its  base  are  surrounded  by  less  intercel- 


8 Contributions  in  Science,  Number  345 


Wake  and  Elias:  Tropical  Salamanders 


lular  matrix  than  are  the  cells  either  proximal  or  distal  to  it. 
This  process  apparently  represents  a primitive  condition  and 
is  reminiscent  of  the  process  seen  in  Ensatina  (Lombard  and 
Wake,  1977),  although  the  process  in  Ensatina  is  less  con- 
tinuous with  the  basibranchial.  The  anterior  basibranchial 
process  is  clearly  a less-derived  homologue  of  the  lingual 
cartilage  that  is  present  in  many  tropical  plethodontids. 

The  vertebral  column  consists  of  relatively  stout  vertebrae, 
with  well-developed  centra.  There  are  one  cervical,  fourteen 
trunk,  one  sacral,  two  caudosacral,  and  a varying  number  of 
caudal  vertebrae  (from  20  to  26  in  individuals  having  com- 
plete tails).  The  atlas  has  no  special  features,  but  the  neural 
arch  is  barely  completed  in  both  cleared  and  stained  adults. 
The  first  trunk  vertebra  is  distinctly  shorter  than  the  re- 
maining vertebrae  and  has  a high,  well-developed  neural 
crest.  Lower  crests  are  present  on  the  next  few  trunk  verte- 
brae. The  centrum  is  large  and  may  be  largely  filled  with 
mineralized  cartilage,  for  only  the  terminal  concavities  are 
clearly  filled  by  unmineralized  intervertebral  cartilage.  All 
trunk  vertebrae  have  well-developed,  separated,  pointed  pro- 
cesses (hyperapophyses)  on  the  posterior  border  of  the  neural 
arch.  The  nerve  route  pattern  is  typical  of  bolitoglossine  sal- 
amanders (Edwards,  1976):  the  first  trunk  vertebra  has  a 
single  foramen  in  front  of  the  transverse  processes,  the  second 
has  single  foramina  both  in  front  of  and  behind  the  transverse 
processes,  and  the  third  and  succeeding  vertebrae  (to  the  end 
of  the  tail)  have  single  foramina  behind  the  transverse  pro- 
cesses. The  transverse  processes  of  the  trunk  vertebrae  are 
well  developed.  The  dorsal  and  ventral  rib  bearers  are  sep- 
arated for  their  entire  lengths.  The  long,  relatively  straight 
processes  extend  beyond  the  lateral  margins  of  the  zygapoph- 
yses.  They  are  sharply  angled  posteriorly  in  the  first  few 
vertebrae  but  are  nearly  perpendicular  to  the  column  over 
most  of  its  length.  The  dorsal  bearer  is  immediately  dorsal 
to  the  ventral  bearer,  but  it  tends  to  be  slightly  more  pos- 
teriorly oriented.  Moderately  long  ribs  with  distinctly  sepa- 
rated heads  are  present  on  all  but  the  last  trunk  vertebra. 
Transverse  processes  of  the  first  caudosacral  vertebra  are 
stout  and  blunt-ended  and  are  swept  somewhat  posteriorly. 
Transverse  processes  of  the  second  caudosacral  vertebra  are 
short,  stout,  and  blunt-ended;  they  are  nearly  perpendicular 
to  the  column  or  are  slightly  anteriorly  oriented.  The  first 
caudal  vertebra  is  short  and  is  associated  with  a distinctly 
constricted  tail  base.  The  relatively  short  transverse  processes 
located  at  the  extreme  anterior  end  of  the  centrum  are  sharply 
oriented  in  an  anterior  direction.  The  transverse  processes 
of  succeeding  caudal  vertebrae  are  progressively  shorter  and 
barely  exceed  the  anterior  zygapophyses  in  length;  they  are 
borne  on  the  extreme  anterior  end  of  each  vertebra,  at  the 
base  of  the  zygapophyses.  The  sharply  tapered  tail  is  deep  as 
a result  of  a thick  glandular  layer  lying  dorsal  to  the  vertebral 
column  but  is  relatively  narrow.  Caudal  vertebrae  generally 
lack  neural  crests,  although  there  may  be  low  and  irregularly 
formed  crests  on  the  first  one  or  two  vertebrae;  however,  the 
caudal  vertebrae  have  very  well-developed  hyperapophyses. 

The  small  hands  and  feet  are  highly  distinctive  (Fig.  4). 
They  are  very  reduced  in  size  and  have  relatively  little  ossified 
tissue.  The  digits  are  fused  to  one  another  for  most  of  their 


lengths  but  are  free  at  their  tips.  Indeed,  the  digits  are  so 
poorly  developed  that  it  is  somewhat  artificial  to  present  a 
phalangeal  formula.  Some  phalanges  are  entirely  cartilagi- 
nous, others  may  have  a tiny  speck  of  ossified  tissue,  and 
only  the  proximal  phalanx  of  the  longest  finger  and  toe  is 
consistently  well  ossified.  An  unusual  feature  is  the  cartilag- 
inous tip  of  many  terminal  phalanges.  In  other  plethodontid 
salamanders,  these  tips  are  well  ossified  and  often  specialized 
in  structure.  The  poor  degree  of  development  suggests  that 
Bradytriton  does  not  use  its  limbs  for  specialized  locomotory 
activity.  A few  individuals  can  be  scored  as  having  a pha- 
langeal formula  of  1,  2,  3,  2,  for  the  hand  and  1,  2,  3,  3,  2, 
for  the  foot,  based  on  the  presence  of  tiny  specks  of  radio- 
opaque material  visible  in  radiographs.  Even  the  metapodial 
elements,  which  are  cylindrical  in  shape,  are  small  and  poorly 
ossified,  and  the  first  digit  of  the  hand  has  an  especially  small 
element.  As  a result  of  the  fusion  of  the  ulnare  and  inter- 
medium, there  are  seven  carpals;  similarly,  there  are  eight 
tarsals  as  a result  of  the  fusion  of  distal  tarsals  4 and  5.  A 
distinct  spur  projects  from  the  shank  of  the  tibia  for  nearly 
its  entire  length,  and  there  is  a low  but  distinct  tibial  crest. 

RANGE.  Bradytriton  silus  is  known  only  from  the  im- 
mediate vicinity  of  the  type  locality  on  the  eastern  slopes  of 
the  Sierra  de  los  Cuchumatanes  in  extreme  northwestern 
Guatemala. 

ETYMOLOGY.  From  silus,  L.  for  “pug-nosed”  referring 
to  the  characteristic  truncated  appearance  of  the  snout. 

GENERIC  SYNOPSES 

Bolitoglossa  Dumeril,  Bibron, 
and  Dumeril 

Oedipus  Tschudi 
Eladinea  Miranda-Ribeiro 
Magnadigita  Taylor 
Palmatotnton  Smith 

TYPE  SPECIES.  Bolitoglossa  mexicana  Dumeril,  Bi- 
bron, and  Dumeril. 

DIAGNOSTIC  CHARACTERS.  Small  to  very  large  sal- 
amanders with  partially  to  fully  webbed  hands  and  feet,  dis- 
tinguished from  members  of  all  other  tropical  salamander 
genera  in  lacking  a sublingual  fold. 

REFERRED  SPECIES.  Bolitoglossa  adspersa  (Peters);  B. 
altamazonica  (Cope);  B.  alvaradoi  Taylor;  B.  arborescandens 
Taylor;  B.  biseriata  Tanner;  B.  borburata  Trapido;  B.  capi- 
tana  Brame  and  Wake;  B.  cerroensis  Taylor;  B.  chica  Brame 
and  Wake;  B.  colonnea  (Dunn);  B.  compacta  Wake,  Brame, 
and  Duellman;  B.  cuchumatana  (Stuart);  B.  cuna  Wake, 
Brame,  and  Duellman;  B.  dofleini  (Werner);  B.  dunni 
(Schmidt);  B.  engelhardti  (Schmidt);  B.  epimela  Wake  and 
Brame;  B.  equatoriana  Brame  and  Wake;  B.  flavimembris 
(Schmidt);  B.  flaviventris  (Schmidt);  B.  franklini  (Schmidt); 
B.  hartwegi  Wake  and  Brame;  B.  helmrichi  (Schmidt);  B. 
hypacra  (Brame  and  Wake);  B.  hgnicolor  ( Peters);  B.  lincolni 
McCoy  and  Walker;  B.  macrinii  (Lafrentz);  B.  marmorea 
(Tanner  and  Brame);  B.  medemi  Brame  and  Wake;  B.  me- 
liana  Wake  and  Lynch;  B.  mexicana  Dumeril,  Bibron  and 


Contributions  in  Science,  Number  345 


Wake  and  Elias:  Tropical  Salamanders  9 


Figure  7.  Feet  of  Chiropterotriton  and  Pseudoeurycea. 

A.  Left  foot  of  an  adult  Chiropterotriton  multidentatus  (39.6  nint  SL)  from  Hidalgo,  Mexico.  Cartilage  is  stippled.  Note  that  distal  tarsal  5 
is  larger  than  distal  tarsal  4 and  articulates  with  the  centrale.  This  arrangement  is  unique  to  Chiropterotriton  within  the  supergenus  Bolitoglossa. 

B.  Left  foot  of  an  adult  Pseudoeurycea  leprosa  (53.9  mm  SL)  from  Veracruz,  Mexico.  Cartilage  is  stippled.  Note  that  distal  tarsal  5 is  smaller 
than  distal  tarsal  4 and  that  it  does  not  articulate  with  the  centrale;  this  is  the  primitive  pattern  in  plethodontid  salamanders.  The  foot  of  this 
species  is  relatively  much  smaller  than  that  of  C.  multidentatus,  for  although  the  feet  as  illustrated  are  about  the  same  size,  the  specimen  of 
Pseudoeurycea  is  much  larger.  Note  that  the  fifth  toe  is  smaller  in  Pseudoeurycea  than  in  Chiropterotriton. 


Dumeril;  B.  minutula  Wake,  Brame,  and  Duellman;  B.  mo- 
no (Cope);  B.  mu/leri  (Brocchi);  B.  nicefori  Brame  and  Wake; 
B.  occidentalis  Taylor;  B.  odonnelli  (Stuart);  B.  orestes  Brame 
and  Wake;  B.  palmata  (Werner);  B.  pandi  Brame  and  Wake; 
B.  peruviana  (Boulenger);  B.  phalarosoma  Wake  and  Brame; 
B.  platydacty/a  (Gray);  B.  ramosi  Brame  and  Wake;  B.  re- 
splendent McCoy  and  Walker;  B.  riletti  Holman;  B.  robusta 
(Cope);  B.  rostrata  (Brocchi);  B.  rufescens  (Cope);  B.  salvinii 
(Gray);  B.  savagei  (Brame  and  Wake);  B.  schizodactyla  Wake 
and  Brame;  B.  schmidti  (Dunn);  B.  silverstonei  Brame  and 
Wake;  B.  sima  (Vaillant);  B.  sooyorum  Vial;  B.  striatula  (No- 
ble); B.  stuarti  Wake  and  Brame;  B.  subpalmata  (Boulenger); 
B.  taylori  Wake,  Brame,  and  Myers;  B.  vallecula  Brame  and 
Wake;  B.  veracrucis  Taylor;  B.  walkeri  Brame  and  Wake;  B. 
yucatana  (Peters). 

RANGE.  Bolitoglossa  has  by  far  the  widest  range  of  any 
of  the  tropical  salamander  genera.  It  occurs  almost  contin- 
uously from  the  lowlands  of  eastern  San  Luis  Potosi,  Mexico, 


in  the  north,  south  to  the  Amazonian  lowlands  of  southern 
Peru,  the  mountains  near  Cochabamba,  Bolivia,  and  south- 
ern Minas  Gerais,  Brazil. 

COMMENT.  Many  species  have  been  added  to  this  large 
genus  during  the  past  two  decades,  but  no  revisionary  study 
has  been  undertaken.  A number  of  undescribed  species  are 
known  to  us.  We  here  place  Bolitoglossa  omniumsanctorum 
in  the  synonymy  of  B.  morio  on  the  basis  of  close  similarity 
of  the  type  specimens  of  B.  omniumsanctorum  to  specimens 
of  B.  morio  in  size,  form,  and  coloration. 

Wake  and  Brame  (1969)  and  Wake  and  Lynch  (1976)  dis- 
cussed a possible  subdivision  of  this  large,  cumbersome  ge- 
nus. Wake  and  Lynch  (1976)  referred  to  “alpha”  and  “beta” 
sister  groups  and  suggested  that  these  might  eventually  be 
recognized  as  separate  genera.  The  beta  group  is  defined  on 
the  basis  of  an  apomorphic  character,  a tail  base  specializa- 
tion (Wake  and  Dresner,  1967).  Members  of  the  alpha  group 
frequently  have  osteological  reductions  and  foot  specializa- 


10  Contributions  in  Science,  Number  345 


Wake  and  Elias:  Tropical  Salamanders 


tions,  but  we  know  of  no  apomorphic  character  or  combi- 
nation of  characters  that  would  unambiguously  define  the 
alpha  group.  These  diagnostic  problems,  combined  with  the 
absence  of  marked  ecological  or  phenetic  differentiation  be- 
tween the  two  groups,  lead  us  to  postpone  formal  division. 

Bradytriton  new  genus 

TYPE  SPECIES.  Bradytriton  si/us  new  species. 

DIAGNOSTIC  CHARACTERS.  This  is  a short-snouted, 
stout-bodied  salamander  with  small  limbs  and  small,  syn- 
dactylous  hands  and  feet.  It  differs  from  all  other  tropical 
salamander  genera  in  having  a laterally  compressed  tail  with 
a well-developed  dorsal  glandular  ridge. 

REFERRED  SPECIES.  Bradytriton  situs  new  species. 

RANGE.  The  unique  species  is  known  only  from  the  im- 
mediate vicinity  of  the  type  locality  on  the  northeastern  slopes 
of  the  Sierra  de  los  Cuchumatanes  in  northwestern  Guate- 
mala. 

Chiropterotriton  Taylor 

TYPE  SPECIES.  Oedipus  multident atus  Taylor. 

DIAGNOSTIC  CHARACTERS.  Small  to  moderately 
large  salamanders  with  relatively  broad,  partially  webbed 
hands  and  feet  and  broad-tipped  fingers  and  toes.  Chirop- 
terotriton differs  from  all  other  tropical  salamander  genera 
in  its  arrangement  of  tarsal  cartilages:  in  Chiropterotriton, 
distal  tarsal  five  is  larger  than  distal  tarsal  four  and  articulates 
with  the  centrale;  in  other  genera,  four  is  larger  than  five, 
and  five  is  excluded  from  articulation  with  the  centrale  (Fig. 
7). 

REFERRED  SPECIES.  Chiropterotriton  arboreus  (Tay- 
lor); C.  chiropterus  (Cope);  C.  chondrostega  (Taylor);  C.  di- 
midiatus  (Taylor);  C.  lavae  (Taylor);  C.  magnipes  Rabb;  C. 
mosaueri  (Woodall);  C.  multidentatus  (Taylor);  C.  prisons 
Rabb. 

RANGE.  The  genus  is  restricted  to  eastern  Mexico,  from 
west-central  Tamaulipas  in  the  north  to  the  mountains  of 
northern  Oaxaca  in  the  south. 

COMMENT.  Most  species  of  this  genus  have  been  stud- 
ied in  some  detail  by  Rabb  (1955,  1958,  1965).  The  major 
unstudied  unit  is  that  comprising  C.  chiropterus  and  C.  lavae. 

Dendrotriton  new  genus 

TYPE  SPECIES.  Oedipus  bromeliacia  Schmidt. 

DIAGNOSIS.  Small,  slender,  long-tailed,  arboreal  sala- 
manders with  long  legs,  broad  hands  and  feet,  and  long, 
broad-tipped  digits.  Dendrotriton  differs  from  Oedipina  by 
its  short  trunk  (14  vs.  18  or  more  trunk  vertebrae)  and  long 
limbs;  from  Nyctanolis  in  having  a single  premaxillary;  from 
Chiropterotriton  in  having  a larger  fourth  than  fifth  tarsal; 
from  Bolitog/ossa  in  possessing  a sublingual  fold;  from  Bra- 
dytriton, Nototriton,  Parvimolge,  Oedipina  and  Thorius  by 
its  lack  of  carpal  and  tarsal  fusions;  from  Lineatriton  by  its 
short  vertebrae  and  long  legs;  and  from  Pseudoeurycea  by  its 
lack  of  prefrontals  and  tibial  spurs. 

REFERRED  SPECIES.  Chiropterotriton  bromeliacia 


(Schmidt);  C.  cuchumatanus  Lynch  and  Wake;  C.  mega- 
rhinus  Rabb;  C.  rabbi  Lynch  and  Wake;  C.  xolocalcae  (Tay- 
lor). 

ETYMOLOGY.  From  dendron,  Gr.  for  tree,  referring  to 
the  arboreal  habits  of  all  known  species,  and  triton,  Gr.,  a 
commonly  used  term  for  salamanders. 

RANGE.  Southwestern  Chiapas,  Mexico,  through  west- 
ern Guatemala. 

COMMENT.  This  group  has  recently  been  studied  in  de- 
tail by  Lynch  and  Wake  (1975),  who  presented  photographs 
of  all  five  species. 

Lineatriton  Tanner 

TYPE  SPECIES.  Spelerpes  lineolus  Cope. 

DIAGNOSTIC  CHARACTERS.  A very  slender,  elon- 
gate, fossorial  form  with  diminutive  limbs,  hands  and  feet, 
and  an  extremely  long  tail.  Lineatriton  is  distinguished  from 
all  other  tropical  salamander  genera  in  having  very  elongated, 
narrow  vertebrae.  Oedipina  has  a similar  body  form  and 
ecology  but  differs  from  Lineatriton  in  having  18  or  more, 
rather  than  14,  trunk  vertebrae. 

REFERRED  SPECIES.  Lineatriton  lineola  (Cope). 

RANGE.  Lineatriton  is  restricted  to  two  disjunct  areas  of 
the  Gulf  slope  of  Mexico:  along  the  lower  slopes  of  the  moun- 
tains of  west-central  Veracruz  and  the  area  around  Los  Tux- 
tlas  in  eastern  Veracruz. 

COMMENT.  This  is  one  of  the  most  distinctive  of  the 
tropical  genera  and  is  apparently  monotypic.  It  has  been  little 
studied  since  the  work  of  Tanner  (1950). 

Nototriton  new  genus 

TYPE  SPECIES.  Spelerpes  picadoi  Stejneger. 

DIAGNOSIS.  Small,  long-tailed  salamanders  of  arboreal, 
terrestrial,  or  semifossorial  habitats.  The  species  have  mod- 
erately long  to  short  legs  and  small  hands  and  feet.  The  in- 
termedium and  ulnare  of  the  manus  and  distal  tarsals  four 
and  five  of  the  pes  are  fused.  The  genus  is  distinguished  from 
other  genera  with  similar  fusions  as  follows:  from  Oedipina 
by  having  only  14  rather  than  18  or  more  trunk  vertebrae; 
from  Bradytriton  by  having  a long,  slender  tail  and  slender 
body;  from  Parvimolge  by  having  the  tail  longer  than  the 
body,  by  having  frontal  processes  fused  together  at  their  base 
(all  species  but  richardi),  and  by  lacking  mesopodia!  and 
hyobranchial  mineralizations;  from  Thorius  by  having  a 
complete  skull  roof  over  the  brain  case;  from  Bo/itoglossa 
by  having  carpal  fusions  and  a sublingual  fold. 

REFERRED  SPECIES.  Chiropterotriton  barbouri 
(Schmidt);  Bo/itoglossa diminuta  Robinson;  Chiropterotriton 
nasalis  (Dunn);  C.  picadoi  (Stejneger);  C.  richardi  (Taylor); 
C.  veraepacis  Lynch  and  Wake. 

ETYMOLOGY.  From  notos,  L.,  referring  to  the  southerly 
distribution  of  the  genus,  and  triton,  L.,  Gr.,  a commonly 
used  term  for  salamanders. 

RANGE.  Nototriton  ranges  from  eastern  Guatemala  to  the 
Meseta  Central  of  Costa  Rica. 

COMMENT.  Most  of  Nototriton  consists  of  what  has  been 


Contributions  in  Science,  Number  345 


Wake  and  Elias:  Tropical  Salamanders  11 


called  the  picadoi  group  (Wake  and  Lynch,  1976)  or  the 
nasa/is  group  (Lynch  and  Wake,  1978)  of  Chiropterotriton. 
The  species  richardi  may  not  be  referable  to  this  genus;  it 
was  originally  described  as  a member  of  the  genus  Parvimolge 
and  has  been  considered  problematic  by  Rabb  (1955),  Wake 
(1966),  Wake  and  Lynch  (1976),  and  Lynch  and  Wake  (1978). 
It  is  more  appropriately  placed  in  Nototriton  than  elsewhere, 
but  it  differs  from  other  members  of  the  genus  in  possessing 
an  unusual  combination  of  ancestral  and  derived  characters 
(Lynch  and  Wake,  1978).  Table  2 was  constructed  by  ignoring 

N.  richardi,  but  we  have  nonetheless  tentatively  assigned  the 
species  to  this  genus  because  it  does  not  come  as  close  to 
fitting  into  any  other  genus.  Further,  we  are  reluctant  to  es- 
tablish a new  genus  for  such  a poorly  known  form. 

The  species  described  as  Bolitoglossa  diminuta  by  Rob- 
inson (1976)  is  also  most  appropriately  placed  in  Nototriton. 
Radiographs  of  the  holotype  reveal  that  it  has  frontal  pro- 
cesses of  the  premaxillary  that  are  fused,  and  it  has  a very 
long  tail  for  its  small  body  size  (53%  of  total  length).  It  was 
reported  to  lack  a sublingual  fold,  and  we  have  been  unable 
to  see  one;  however,  the  holotype,  a mature  female,  is  very 
small  (31.1  mm  SL),  and  it  may  be  impossible  to  determine 
the  status  of  this  character  without  histological  sections.  No- 
totriton richardi  has  a very  reduced  sublingual  fold.  It  was 
on  the  basis  of  the  apparent  absence  of  this  fold  in  the  ho- 
lotype of  B.  diminuta  that  the  senior  author  urged  Robinson 
to  describe  the  species  as  a Bolitoglossa,  even  though  Rob- 
inson had  originally  intended  to  place  it  in  the  genus  Chi- 
ropterotriton. 

Nyctano/is  Elias  and  Wake 

TYPE  SPECIES.  Nyctanolis  pernix  Elias  and  Wake. 

DIAGNOSTIC  CHARACTERS.  This  is  a large,  long- 
legged,  long-fingered,  long-toed,  long-tailed  taxon  that  differs 
from  all  other  tropical  salamander  genera  in  having  paired 
premaxillary  bones. 

REFERRED  SPECIES.  Nyctanolis  pernix  Elias  and 
Wake. 

RANGE.  The  unique  species  is  known  from  the  north- 
eastern slopes  of  the  Sierra  de  los  Cuchumatanes  in  NW 
Guatemala;  near  the  Lagos  de  Montebello  in  nearby  Chiapas, 
Mexico;  and  from  the  Sierra  de  las  Minas  in  eastern  Gua- 
temala. 

Oedipina  Keferstein 

Oedipina  Keferstein 
Ophiobatrachus  Gray 
? Haptoglossa  Cope 
Oedopinola  Hilton 

TYPE  SPECIES.  Oedipina  uniformis  Keferstein. 

DIAGNOSTIC  CHARACTERS.  Slender,  elongate  sala- 
manders of  moderate  to  large  size  with  very  small  limbs, 
hands,  and  feet,  and  a very  long  tail.  Oedipina  is  distinguished 
from  all  other  tropical  salamander  genera,  including  Linea- 
triton,  the  only  genus  that  resembles  it  externally,  by  having 
18  to  22  (rather  than  14)  trunk  vertebrae. 


REFERRED  SPECIES.  Oedipina  alfaroi  Dunn;  O.  al- 
tura  Brame;  O.  carab/anca  Brame;  O.  col/aris  (Stejneger);  O. 
complex (D\inn)\  O.  cyclocauda  Taylor;  O.  e/ongata  (Schmidt); 

O.  grandis  Brame  and  Duellman;  O.  ignea  Stuart;  O.  par- 
vipes  (Peters);  O.  paucidentata  Brame;  O.  poelzi  Brame;  O. 
pseudouniformis  Brame;  O.  stuarti  Brame;  O.  taylori  Stuart; 

O.  uniformis  Keferstein. 

RANGE.  Oedipina  ranges  from  south-central  Chiapas, 
Mexico,  through  all  of  Central  America  and  western  Colom- 
bia to  northwestern  Ecuador. 

COMMENT.  This  is  an  exceptionally  well-defined  genus. 
Since  the  revision  by  Brame  (1968),  only  one  additional 
species,  O.  grandis,  has  been  described,  and  there  have  been 
no  other  taxonomic  changes. 

Parvimolge  Taylor 

TYPE  SPECIES.  Oedipus  townsendi  Dunn. 

DIAGNOSTIC  CHARACTERS.  Diminutive  salaman- 
ders with  relatively  stocky  bodies,  short  legs  with  small  hands 
and  feet,  and  short,  stout  tails.  It  is  distinguished  from  other 
genera  with  small  species  as  follows:  from  Bolitoglossa  in 
having  a sublingual  fold;  from  Dendrotriton  and  Chiroptero- 
triton in  having  distal  tarsals  four  and  five  fused  and  the 
intermedium  and  ulnare  fused;  from  Nototriton  in  having  a 
short  tail  and  lacking  fused  frontal  processes  of  the  premax- 
illary; from  Thorius  in  having  a complete  skull  roof  over  the 
braincase. 

REFERRED  SPECIES.  Parvimolge  townsendi  (Dunn). 

COMMENT.  Two  other  species,  praecellens  and  rich- 
ardi, have  been  included  in  the  genus.  Neither  seems  to  be 
closely  related  to  townsendi,  and  we  here  refer  them  to  the 
genera  Pseudoeurycea  and  Nototriton,  respectively. 

Pseudoeurycea  Taylor 

TYPE  SPECIES.  Spelerpes  leprosus  Cope. 

DIAGNOSTIC  CHARACTERS.  A diverse  group  of 
moderate-sized  to  very  large  salamanders  with  moderate  to 
long  limbs,  well-developed  hands  and  feet  in  which  the  mid- 
dle digits  are  markedly  longer  than  the  outer  ones  (Fig.  7), 
and  a tail  about  as  long  as  the  body.  The  species  are  very 
generalized  in  morphology.  The  genus  differs  from  Nyctanolis 
in  having  a single  premaxillary  and  shorter  limbs  and  tail; 
from  Dendrotriton  in  having  prefrontals  and  tibial  spurs; 
from  Bolitoglossa  in  having  a sublingual  fold;  from  Chirop- 
terotriton in  having  a fourth  distal  tarsal  that  articulates  with 
the  fibulare  and  is  larger  than  the  fifth  (Fig.  7);  from  all  other 
genera  in  having  separate  distal  tarsals  four  and  five. 

REFERRED  SPECIES.  Pseudoeurycea  altamontana 
(Taylor);  P.  anitae  Bogert;  P.  bellii  (Gray);  P.  brunnata 
Bumzahem  and  Smith;  P.  cephalica  (Cope);  P.  cochranae 
(Taylor);  P.  conanti  Bogert;  P.  exspectata  Stuart;  P.firscheini 
Shannon  and  Werler;  P.  gadovii  (Dunn);  P.  ga/eanae  (Tay- 
lor); P.  goebeli  (Schmidt);  P.juarezi  Regal;  P.  leprosa  (Cope); 

P.  melanomolga  (Taylor);  P.  mystax  Bogert;  P.  nigroma- 
culata  (Taylor);  P.  praecellens  (Rabb);  P.  rex  (Dunn);  P.  rob- 
ertsi  (Taylor);  P.  scandens  Walker;  P.  smithi  (Taylor);  P. 
unguidentis  Smith  and  Taylor;  P.  werleri  Darling  and  Smith. 


12  Contributions  in  Science,  Number  345 


Wake  and  Elias:  Tropical  Salamanders 


RANGE.  The  genus  ranges  from  eastern  Sonora,  Mexico, 
in  the  northwest,  and  west-central  Tamaulipas,  Mexico,  in 
the  northeast,  through  most  of  eastern  and  south-central 
Mexico  into  western  and  southern  Guatemala. 

COMMENT.  Few  characters  bind  this  assemblage  ofgen- 
eralized  species  together.  The  genus  lacks  any  autapomor- 
phies  and  is  mainly  defined  by  exclusion.  A few  species  stand 
out  from  the  rest  as  distinct  morphological  entities  (e.g.,  P. 
bellii,  P.  praecel/ens,  P.  werleri).  The  gadovii,  rex,  and  leprosa 
groups  are  the  core  of  the  genus  and  appear  to  be  closely 
interrelated  (Maxson  and  Wake,  1981).  We  here  assign  prae- 
cel/ens to  Pseudoeurycea.  Rabb  (1955)  placed  it  in  the  genus 
Parvimolge  with  some  hesitation.  It  differs  from  P.  townsendi 
in  that  it  is  larger  and  stouter  and  has  prefrontal  bones.  Rabb 
(1955)  reported  that  P.  praecel/ens  had  some  mineralization 
of  the  hyobranchial  apparatus,  as  in  P.  townsendi ; however, 
we  have  been  unable  to  see  this  feature  in  our  radiographs 
of  the  unique  holotype.  The  species  fits  reasonably  well  in 
Pseudoeurycea,  and  Rabb  suggested  that  it  might  prove  to 
be  a relative  of  P.  cephalica. 

Thor ius  Cope 

TYPE  SPECIES.  Thorius  pennatulus  Cope. 

DIAGNOSTIC  CHARACTERS.  Diminutive,  slender 
salamanders  with  short  limbs  and  small  hands  and  feet.  Tho- 
rius is  distinguished  from  all  other  tropical  salamanders  by 
two  characters:  the  suborbital  groove  intersects  the  lip,  and 
the  skull  is  incompletely  ossified,  leaving  the  braincase  un- 
covered by  bone  (the  parietal  bones  have  an  especially  large 
gap  separating  them). 

REFERRED  SPECIES.  Thorius  duhitus  Taylor;  T.  mac- 
dougalli  Taylor;  T.  maxil/abrochus  Gehlbach;  T.  minutissi- 
mus  Taylor;  T.  narisovalis  Taylor;  T.  pennatulus  Cope;  T. 
pulmonaris  Taylor;  T.  schmidti  Gehlbach;  T.  troglodytes 
Taylor. 

RANGE.  The  genus  ranges  from  the  eastern  margins  of 
the  Mexican  Plateau  into  the  mountains  of  northern  and 
central  Oaxaca,  and  in  the  Sierra  Madre  del  Sur  of  southern 
Oaxaca  and  Guerrero,  Mexico. 

COMMENT.  The  genus  has  recently  been  studied  in  de- 
tail by  Freeman  (1977)  and  Hanken  (1980),  but  the  taxo- 
nomic results  of  these  unpublished  theses  are  not  yet  avail- 
able. All  of  the  described  species  (except  perhaps  T. 
maxillabrochus)  are  valid,  but  there  are  numbers  of  addi- 
tional undescribed  species. 

DISCUSSION 

The  supergenus  Bolitoglossa  was  redefined  by  Elias  and  Wake 
(1983)  on  the  basis  of  hyobranchial  and  tail  base  character- 
istics. All  genera  considered  here  have  the  features  of  the 
supergenus.  Our  main  departure  from  Wake  (1966)  is  that 
two  formerly  diagnostic  features  are  deemphasized.  First, 
Nyctanolis  has  two  premaxillary  bones;  thus,  the  supergenus 
Bolitoglossa  can  no  longer  be  characterized  by  the  presence 
of  a single,  fused  bone.  Second,  Nyctanolis  and  some  species 
of  Pseudoeurycea  have  a lateral  parietal  spur  that  is  only 


poorly  developed:  in  Nyctanolis  the  spur  may  be  little  more 
than  a low  ridge  on  the  underside  of  the  bone,  and  in  some 
Pseudoeurycea  the  “spur”  is  really  only  a small  lateral  lobe. 
Still,  at  least  some  form  of  parietal  “spur”  is  present  in  all 
members  of  the  supergenera  Batrachoseps  and  Bolitoglossa. 

With  the  new  genera,  the  supergenus  Bolitoglossa  includes 
1 1 genera.  Nyctanolis,  Bradytriton,  Parvimolge,  and  Linea- 
triton  are  monotypic;  the  other  genera  include  from  five  (Den- 
drotriton)  to  over  sixty  (Bolitoglossa)  species.  Additional  un- 
described species  that  are  known  to  us  will  increase  the  number 
of  species  in  such  large  genera  as  Bolitoglossa,  Pseudoeurycea, 
and  Thorius  but  will  not  affect  the  monotypic  genera. 

The  genera  Bolitoglossa,  Lineatriton,  Oedipina,  Pseudoeu- 
rycea, and  Thorius  are  essentially  unchanged  by  our  analysis. 
The  content  of  Parvimolge  is  reduced  to  a single  species  by 
transfer  of  the  species  praecel/ens  to  Pseudoeurycea.  The  ge- 
nus most  strongly  affected  is  Chiropterotriton.  As  a result  of 
the  shift  of  the  old  bromeliacia  species  group  (Lynch  and 
Wake,  1975)  to  the  new  genus  Dendrotriton  and  of  the  old 
picadoi( Wake  and  Lynch,  1 976)  or  nasalis  (Lynch  and  Wake, 

1 978)  species  group  to  the  new  genus  Nototriton,  Chiroptero- 
triton has  been  reduced  to  a group  of  nine  species  that  inhabit 
eastern  Mexico.  The  species  formerly  known  as  Bolitoglossa 
diminuta  is  included  in  Nototriton.  The  other  two  new  genera, 
Nyctanolis  and  Bradytriton,  are  based  on  newly  discovered 
species. 

Hecht  and  Edwards  (1976a,  1 976b)  and  Hecht  (1976)  dis- 
cuss the  relative  information  content  of  different  types  of 
shared  derived  character  states  (synapomorphies)  for  the  re- 
construction of  phylogeny.  Ordered  by  increasing  significance 
(basically,  the  degree  of  confidence  one  has  in  their  inter- 
pretation) in  phylogenetic  reconstruction  are  five  classes  of 
synapomorphies: 

“(I)  Shared  and  derived  character  states  which  are  the  result 
of  loss.  Such  loss  characters  are  the  zero  state  because 
there  is  no  developmental  evidence. 

(II)  Shared  and  derived  character  states  which  are  the  results 
of  simplification  or  reduction  of  complex  structures  as 
indicated  by  comparative  or  developmental  anatomy. 

(III)  Shared  and  derived  character  states  which  are  the  result 
of  growth  and  developmental  processes  dependent  on 
size,  age,  and  hormonal  and  other  physiological  rela- 
tionships. Characters  related  to  allometric  functions  are 
the  best  examples. 

(IV)  Shared  and  derived  character  states  which  are  highly 
integrated  functionally  and  are  subject  to  directional  se- 
lection. 

(V)  Shared  and  derived  character  states  which  are  unique 
and  innovative  in  structure  (the  most  important  type  of 
information).”  (Lrom  Hecht  and  Edwards,  1976a.) 

Our  analysis  in  this  paper  is  based  on  the  18  characters  out- 
lined earlier.  The  apomorphic  characters  we  have  chosen 
include  seven  that  are  autapomorphic  (Table  1).  These  are 
not  useful  in  phylogenetic  analysis  but  are  included  because 
they  aid  in  diagnosing  genera.  The  1 1 remaining  characters 
vary  in  degree  of  utility.  Six  involve  morphological  trans- 
formation, and  five  involve  losses  of  elements.  We  interpret 


Contributions  in  Science,  Number  345 


Wake  and  Elias:  Tropical  Salamanders  13 


our  loss  characters  as  falling  into  the  third  category  of  Hecht 
and  Edwards  (1976)  rather  than  the  first,  for  we  have  some 
ontogenetic  information  concerning  the  way  in  which  four 
of  the  five  characters  are  lost.  We  believe  that  loss  of  septo- 
maxillaries,  prefrontals,  preorbital  processes  of  vomers,  and 
columellae  (or  stylar  processes  of  the  opercula)  are  all  pae- 
domorphic  characters  (Lynch  and  Wake,  1978;  Alberch  et 
al.,  1979;  Alberch  and  Alberch,  1981;  Elias,  in  press),  but 
whether  they  should  all  be  counted  as  independent  characters 
or  considered  to  be  parts  of  a single  “global”  paedomorphic 
process  is  unresolved. 

The  six  remaining  characters  ( 1,  3,  4,  5,  10,  1 1 ) all  involve 
transformations.  Characters  10  and  1 1 relate  to  ontogenetic 
phenomena  and  fall  into  Hecht  and  Edwards's  third  category. 
Transformation  characters  1,  3,  4,  and  5 are  parts  of  inte- 
grated, functionally  significant  systems;  they  fall  in  the  fourth 
category  of  Hecht  and  Edwards. 

Other  authors  have  used  additional  characters  in  generic 
diagnoses  of  tropical  salamanders.  Some  of  these  characters 
(e.g.,  a posteriorly  directed  spur  on  the  squamosal  in  Thorius) 
are  autapomorphies  and  of  little  use  in  phylogenetic  analysis. 
Others  occur  in  all  members  of  one  genus  but  also  appear 
irregularly  in  one  or  a few  species  in  some  other  genera  (e.g., 
the  complex  tail  base  present  in  some  Pseudoeurycea).  Other 
characters,  such  as  the  shape  and  degree  of  ossification  of  the 
intervertebral  articulation  (used  by  Taylor,  1 944),  are  subject 
to  extreme  ontogenetic  variation  (Wake,  1 970),  and  we  have 
not  had  sufficient  material  to  perform  a complete  analysis. 
Still  other  characters,  such  as  external  shape  of  hands  and 
feet,  shape  of  terminal  digits,  length  of  ribs,  and  fusion  or 
nonfusion  of  transverse  processes,  have  proven  impossible 
to  codify  consistently  for  all  neotropical  species,  but  these 
may  well  have  utility  for  liner  analysis  of  intra-  and  inter- 
generic relationships  in  the  future.  The  hyobranchial  appa- 
ratus (features  of  the  tip  of  the  first  basibranchial,  including 
length  and  shape  of  radii  and  detailed  structure  of  the  lingual 
cartilage  and  associated  musculature)  holds  much  promise 
as  a source  of  additional  characters  (Tanner,  1952;  Lombard 
and  Wake,  1977).  However,  this  structure  requires  detailed 
histological  study,  and  we  have  examined  too  few  species  to 
discern  broad  patterns.  Future  studies  should  involve  com- 
parative ontogenetic  analyses,  for  heterochronic  modes  of 
evolution  have  been  strongly  implicated  in  salamanders 
(Wake,  1966;  Alberch  et  al.,  1979;  Alberch  and  Alberch, 
1981). 

Biochemical,  immunological,  and  cytological  characters  are 
useful  in  analyzing  phylogenetic  relationships  in  the  super- 
genus Bo/itoglossa.  Recently  Maxson  and  Wake  (1981)  used 
immunological  approaches  to  study  albumin  evolution  in 
Pseudoeurycea  and  Chiropterotriton  (sensu  lato).  Extensive 
evolution  of  albumin  has  taken  place  in  these  groups,  and 
immunological  methods  will  be  most  useful  at  the  intrage- 
neric level  of  comparison.  In  reference  to  the  present  generic 
review,  the  principal  result  of  interest  from  the  immunolog- 
ical work  is  the  finding  that  Chiropterotriton  (antisera  from 
C.  multidentatus)  and  Dendrotriton  (antisera  from  D.  bro- 
meliacia)  are  more  similar  to  Pseudoeurycea  (antisera  from 
five  species)  than  they  are  to  each  other.  This  finding  cor- 


roborates our  conclusion  that  the  two  genera  are  distinct.  No 
antisera  were  prepared  for  species  of  Nototriton,  but  one-way 
tests  were  made  to  some  species  of  the  genus.  Immunological 
distances  between  species  of  Chiropterotriton  and  Dendro- 
triton are  about  equivalent  to  those  between  Chiropterotriton 
and  Nototriton,  but  immunological  distances  between  Den- 
drotriton and  Nototriton  are  considerably  less.  Distances 
within  Dendrotriton  (maximum  21)  are  much  less  than  be- 
tween Dendrotriton  and  Nototriton  (minimum  64). 

Some  data  on  chromosomes  are  available  (Leon  and  Kezer, 
1978;  J.  Kezer  and  S.  Sessions,  pers.  comm.).  All  members 
of  the  supergenus  Bo/itoglossa  so  far  examined  have  a haploid 
chromosome  number  of  13,  as  do  members  of  the  genus 
Batrachoseps.  All  other  plethodontids  examined  have  a hap- 
loid number  of  14.  The  species  of  Dendrotriton  (D.  brome- 
liacia,  D.  cuchumatanus,  D.  rabbi ) and  Nototriton  ( N . 
picadoi,  N.  veraepacis)  so  far  examined  have  a distinct  het- 
eromorphism (presumably  XY)  in  the  thirteenth  pair  of  chro- 
mosomes. This  pattern  has  been  found  in  some  species  of 
Thorius  and  Oedipina  but  not  in  Chiropterotriton,  Pseu- 
doeurycea, Lineatriton,  Bo/itoglossa,  or  Parvimo/ge  (chro- 
mosomes of  Nyctanolis  and  Bradytriton  have  not  been  ex- 
amined). These  data  lend  further  support  to  our  decision  to 
separate  Dendrotriton  and  Nototriton  from  Chiropterotriton. 

From  the  1 8 characters  analyzed  we  are  unable  to  conclu- 
sively demonstrate  the  monophyletic  status  of  3 of  the  1 1 
bolitoglossine  genera:  Dendrotriton,  Nototriton,  and  Pseu- 
doeurycea (Table  1 ).  A genus  appearing  in  Table  1 must  have 
at  least  one  apomorphic  character  (a  synapomorphy  for  the 
species  of  the  genus)  that  is  not  shared  with  any  other  given 
genus  in  order  for  us  to  consider  it  to  be  monophyletic. 

All  of  the  apomorphies  shown  for  Dendrotriton  in  Table 
1 are  shared  with  Oedipina ; nevertheless,  in  other  characters, 
which  proved  impossible  to  score  for  all  neotropical  genera, 
Dendrotriton  shows  derived  states  not  found  in  Oedipina. 
Species  of  Dendrotriton,  all  of  which  are  arboreal,  show  such 
derived  features  as  relatively  long  legs,  with  long,  well-de- 
veloped digits  and  expanded  terminal  phalanges.  Oedipina, 
all  of  which  are  semi-  to  fully  fossorial,  have  very  short  legs, 
with  short,  erratically  developed  digits  and  unspecialized 
phalanges.  The  limb  states  of  the  two  genera  are  derived  in 
opposite  directions  from  the  ancestral  state.  Thus,  despite 
the  absence  of  defining  synapomorphies  in  our  formal  tab- 
ulation, we  consider  Dendrotriton  to  be  a well-defined,  mono- 
phyletic assemblage. 

The  situation  with  Pseudoeurycea  is  far  more  complex. 
This  genus  is  something  of  a “waste-basket,”  notwithstand- 
ing arguments  of  Baird  (1951)  to  the  contrary.  It  has  only  a 
single  apomorphy  (fused  premaxillaries,  character  1 1,  Table 
1 ),  and  this  is  shared  with  all  genera  of  the  supergenus  except 
Nyctanolis.  All  of  the  other  genera  are  easily  distinguished 
from  Pseudoeurycea  on  the  basis  of  a variety  of  synapo- 
morphies and  autapomorphies.  The  species  of  Pseudoeurycea 
are  relatively  diverse  in  external  structure  and  behavior  but 
are  basically  similar  in  their  generalized  osteology,  such  that 
there  is  no  clear  osteological  basis  for  subdividing  the  genus 
into  monophyletic  assemblages.  The  generotype  is  P.  leprosa; 
it  and  some  allied  species  have  a plesiomorphic  character 


14  Contributions  in  Science,  Number  345 


Wake  and  Elias:  Tropical  Salamanders 


(septomaxillary  bones)  and  an  apomorphic  character  (a  com- 
plex tail  base,  not  coded  here,  but  rather  similar  to  the  tail 
base  seen  in  Bolitog/ossa  beta.  Wake  and  Lynch,  1976).  It 
may  eventually  be  possible  to  formally  recognize  this  group 
taxonomically,  but  the  remaining  species  in  the  present  genus 
Pseudoeurycea  would  still  constitute  a very  disparate  assem- 
blage. Maxson  and  Wake  (1981)  have  shown  that  species  of 
Pseudoeurycea  are  greatly  divergent  in  albumin  structure. 
One  group  that  is  not  well  defined  morphologically  can  be 
defined  reasonably  well  on  a biochemical  basis  (the  combined 
gadovii  and  rex  groups  of  Wake  and  Lynch,  1976,  together 
with  some  members  of  other  groups).  However,  an  electro- 
phoretic analysis  in  progress  (Yang  and  Wake,  unpublished 
data)  discloses  that  most  species  are  very  distinct  from  one 
another  and  that  groups  of  species  are  not  readily  apparent. 
Pseudoeurycea  may  be  a paraphyletic  taxon,  the  members  of 
which  have  retained  relatively  conservative  morphologies. 
In  this  respect,  the  genus  presents  a taxonomic  problem  sim- 
ilar to  that  encountered  in  the  genus  Plethodon  (Larson  et 
al.,  1981),  another  taxon  based  largely  on  plesiomorphic 
characteristics.  We  do  not  think  that  Pseudoeurycea,  as  pres- 
ently constituted,  represents  the  remnants  of  the  ancestral 
stock  of  all  tropical  salamanders,  for  in  many  respects  mem- 
bers of  Nyctanolis  and  Chiropterotriton  are  more  generalized 
osteologically.  For  example,  C.  priscus  has  relatively  large, 
well -developed  septomaxillary  bones,  primitive  elements  that 
occur  only  occasionally  in  the  few  species  of  Pseudoeurycea 
that  have  them.  Thus,  our  character  scorings  must  be  used 
cautiously  when  making  phylogenetic  interpretations. 

Notototnton,  although  it  is  readily  distinguished  from  most 
genera,  has  no  apomorphies  that  are  not  shared  with  Oedi- 
pina.  This  situation  is  superficially  similar  to  the  problem  of 
Dendrotriton  versus  Oedipina,  but  the  three  synapomorphies 
shared  by  Nototriton  and  Oedipina  are  a different  set.  Fur- 
ther, unlike  Dendrotriton,  Nototriton  may  be  a paraphyletic 
group  that  includes  lineages  of  a pr e-Oedipina  stock  (see 
Lynch  and  Wake,  1978).  For  example,  N.  richardi,  a poorly 
known  species  of  which  fewer  than  ten  specimens  have  been 
collected,  appears  to  be  more  primitive  than  other  Nototriton 
in  some  respects  but  more  derived  in  others.  It  could  be 
placed  in  Oedipina,  but  it  lacks  an  increased  number  of  trunk 
vertebrae  (character  1 7),  the  synapomorphy  that  most  strong- 
ly distinguishes  the  species  of  Oedipina  from  all  other  tropical 
plethodontids.  The  remaining  species  of  Nototriton  form  a 
well-defined  assemblage  (see  Lynch  and  Wake,  1978).  As 
recognized  here,  Nototriton  is  a diverse  group  consisting  of 
several  rare,  secretive,  and  poorly  known  species.  At  this 
time  we  are  unable  either  to  define  autapomorphies  for  the 
genus  or  to  subdivide  it  into  clearly  monophyletic  sublin- 
eages.  By  joining  Oedipina  and  Nototriton  to  form  a single 
genus,  we  could  claim  monophyly  for  the  group  as  a whole, 
but  this  would  severely  blur  the  limits  of  the  present  genus 
Oedipina,  one  of  the  most  tightly  knit,  well-diagnosed,  and 
highly  apomorphic  genera  of  salamanders.  Species  of  Noto- 
triton share  a number  of  structural  and  ecological  similarities, 
and  they  are  sharply  distinguished  from  Oedipina  by  the 
synapomorphies  of  the  latter  group.  Accordingly,  we  choose 
to  recognize  both  Oedipina  and  Nototriton,  although  the  lat- 


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supergenus 

Bolitoglossa 

Figure  8.  Maximum  parsimony  cladogram  of  genera  in  the  tribe 
Bolitoglossini.  The  numbered  characters  (Table  1 ) are  present  in 
their  derived  condition  in  all  taxa  above  the  point  in  the  dendrogram 
where  the  lines  occur.  Reversals  are  circled.  The  tree  treats  character 
9 as  if  it  were  a synapomorphy  in  its  reversed  form  (i.e.,  resembling 
the  presumed  primitive  condition).  Parallelisms  are  not  illustrated. 


ter  genus  may  ultimately  not  prove  to  be  a monophyletic 
assemblage. 

We  used  the  data  in  Table  1 to  compute  Wagner  trees, 
using  the  method  of  cladistic  inference  by  parsimony  (Farris, 
1970),  but  with  one  modification.  We  fixed  the  root  of  the 
tree  a priori  by  declaring  Nyctanolis  to  be  the  first  derivative 
branch.  We  believe  that  the  premaxillary  character  is  con- 
siderably more  substantial  than  any  other  in  the  matrix  (be- 
cause it  is  conservative  and  relatively  well  understood;  see 
Wake,  1966),  and  we  accordingly  used  the  rooting  method 
to  weight  this  character  heavily.  The  resulting  tree  is  pre- 
sented in  Figure  8. 

There  are  reasons  for  questioning  the  validity  of  this  max- 
imum parsimony  tree.  Although  there  are  six  stems  char- 
acterized by  synapomorphies,  two  of  these  are  reversed  later 
in  the  dendrogram.  Only  three  of  the  six  synapomorphic 


Contributions  in  Science,  Number  345 


Wake  and  Elias:  Tropical  Salamanders  15 


characters  (3,  4,  5)  are  “high-quality”  characters,  i.e.,  char- 
acters that  fall  into  Hecht  and  Edwards’s  (1976a)  category  4 
and  constitute  parts  of  integrated  systems.  The  significance 
of  character  5 is  unclear;  the  long,  tubular  mental  glands  of 
certain  of  the  genera  (e.g.,  Oedipina,  Bradytriton ) are  unusual 
but  may  represent  extreme  cases  of  subtler  variation  that 
would  require  histological  documentation.  Note  that  the  den- 
drogram requires  a reversal  of  this  character  in  the  stem 
leading  to  Thorius.  It  seems  more  likely  that  tubular  glands 
have  evolved  in  parallel,  a hypothesis  that  is  supported  by 
the  observation  that  some  species  of  the  unrelated  North 
American  genus  Eurycea  have  such  glands.  Characters  3 and 
4 involve  fusions  of  carpals  and  tarsals.  Although  these  char- 
acters may  be  more  complex  than  those  that  involve  loss  of 
elements,  they  nevertheless  require  little  more  than  simple 
developmental  modifications.  We  know  that  character  3 has 
evolved  independently  in  extra-tropical  plethodontids  (e.g.. 
Amides  hardii and  some  species  of  Batrachoseps\  Wake,  1 966). 
However,  among  plethodontids  character  4 (tarsal  fusions) 
appears  to  be  unique  to  the  genera  reported  here.  It  is  possible 
that  distal  tarsals  4 and  5 are  fused  in  Balrachoseps,  rather 
than  5 being  lost  as  Wake  (1966)  has  suggested. 

Because  we  have  fixed  the  root  of  the  tree  with  Nyctanolis, 
character  1 1 will  be  a common  feature  of  most  possible  den- 
drograms. Characters  6 and  7 fall  into  Hecht  and  Edwards’s 
third  weighting  category,  for  we  have  some  ontogenetic  in- 
formation concerning  them  (unpublished  and  Elias,  in  press). 
We  think  that  both  characters  6 and  7 reflect  parallel  evo- 
lution, for  both  characters  refer  to  features  that  appear  rel- 
atively late  in  the  development  of  related  species.  Character 
7 (loss  of  preorbital  processes  of  the  vomer)  has  evolved 
independently  in  some  species  of  Balrachoseps  (Marlow  et 
al.,  1979).  Scoring  for  character  6 (loss  of  stylus  of  the  oper- 
culum) requires  a certain  degree  of  subjectivity,  because  the 
process,  where  it  appears  at  all,  is  invariably  very  small  in 
members  of  the  supergenus  Bolitoglossa. 

The  final  significant  character  in  Figure  8 is  the  presence 
or  absence  of  septomaxillary  bones  (character  9).  Most  mem- 
bers of  the  supergenus  Bolitoglossa  lack  septomaxillaries,  and 
a questionable  feature  of  this  tree  is  that  it  contains  a stem 
for  which  the  synapomorphy  is  a postulated  reversal  (reap- 
pearance of  septomaxillaries).  This  reversal  unites  Pseu- 
doeurvcea,  Chiropterotriton.  and  Dendrotriton,  three  genera 
that  share  no  additional  synapomorphies  and  have  little  else 
in  common.  Because  the  first  two  of  these  genera  are  exten- 
sively plesiomorphic,  no  system  of  analysis  can  deal  with 
them  easily. 

Septomaxillary  bones  are  not  well  developed  in  any  trop- 
ical salamander.  They  appear  regularly  in  some  Chiroptero- 
triton (Rabb,  1956;  Wake,  1966)  but  are  absent  in  others 
(e.g.,  C.  dimidiatus).  Septomaxillary  bones  are  extremely  small 
and  variably  present  in  four  of  the  five  species  of  Dendro- 
triton (Lynch  and  Wake,  1975,  1978).  Septomaxillaries  also 
occur  in  at  least  some  individuals  of  several  species  of  Pseu- 
doeurycea  (P.  cephalica,  P.  leprosa,  P.  werleri,  and  P.  nigro- 
maculata)  and  occasionally  in  other  bolitoglossine  genera. 
Wake  (1966)  reports  their  presence  in  individuals  of  Parvi- 
molge  townsendi,  Oedipina  uniformis,  Thorius  pennatulus, 


and  three  species  of  Bolitoglossa.  We  hypothesize  that  trop- 
ical salamanders  as  a group  have  undergone  osteological  re- 
duction, and  that  such  reduction  may  have  characterized  the 
ancestral  stock.  Loss  of  the  septomaxillaries  can  be  inter- 
preted as  a manifestation  of  a general  paedomorphic  trend 
in  the  group  (Wake,  1966;  Alberch  and  Alberch,  1981).  Be- 
cause these  bones  are  so  well  developed  in  Chiropterotriton 
relative  to  other  members  of  the  supergenus  Bolitoglossa.  we 
consider  it  unlikely  that  a reversal  of  this  character  (i.e.,  re- 
evolution of  septomaxillaries  following  their  loss  in  an  ances- 
tor) has  occurred  in  the  genus.  Chiropterotriton  is  the  only 
tropical  genus  in  which  the  presence  of  septomaxillaries  is 
universal  in  adults  of  most  species  (all  but  the  paedomorphic 
C.  dimidiatus).  The  loss  of  septomaxillaries  in  certain  Chi- 
ropterotriton species  parallels  the  loss  of  the  bones  in  other 
genera,  and  the  same  underlying  mechanism  could  be  in- 
volved in  all  cases. 

Paedomorphosis  is  not  necessarily  a unidirectional  pro- 
cess, and  some  reversals  are  to  be  expected.  Thus,  septo- 
maxillaries may  have  reappeared  in  Dendrotriton  and  Pseu- 
doeurycea.  following  their  loss  in  the  respective  ancestral 
stocks.  The  bones  occur  irregularly  even  in  those  species 
where  they  are  found  at  all.  The  rare  appearance  of  septo- 
maxillaries in  other  genera  is  likely  to  be  atavistic,  i.e.,  based 
on  a reversal  of  an  original  paedomorphic  event. 

Summarizing  our  arguments,  linkage  of  Chiropterotriton. 
Dendrotriton.  and  Pseudoeurycea  on  the  basis  of  a postulated 
shared  reversal  of  a single  character  is  not  warranted.  If  the 
septomaxillary  character  is  omitted  from  the  Wagner  anal- 
ysis, the  resulting  dendrogram  includes  a stem  uniting  these 
same  three  genera.  This  time  the  stem  is  based  on  another 
shared  reversal,  separation  of  tarsal  elements  (character  4). 
We  consider  such  a reversal  to  be  unlikely  but  not  impossible. 
Instances  are  known  in  which  species  with  fused  tarsals  show 
polymorphism  for  the  trait,  so  reversals  could  occur  (Wake, 
1966;  Wake  and  Brame,  1969;  Alberch  and  Alberch,  1981). 
We  conclude  that  parsimony  methods  must  be  used  cau- 
tiously where  extensive  parallelism  pervades  an  evolutionary 
radiation. 

An  alternative  method  of  analyzing  our  data  is  illustrated 
in  Figure  9.  This  dendrogram  is  based  on  the  zoogeographic 
and  phyletic  arguments  of  Wake  and  Lynch  (1976,  fig.  32), 
who  recognized  a "Mexican  Group”  of  genera  including 
Pseudoeurycea.  Chiropterotriton  ( sensu  stricto ),  Parvimolge, 
Lineatriton.  and  Thorius.  Using  the  present  suite  of  osteo- 
logical characters,  we  can  find  no  justification  for  recognition 
of  a clade  including  these  five  genera;  so  we  have  indicated 
a five-way  division  separating  them  at  the  second  level  of 
the  dendrogram  and  grouping  them  to  the  left  of  the  figure. 
Although  this  dendrogram  has  relatively  little  internal  struc- 
ture, it  does  distinguish  two  groups  of  three  and  four  genera, 
respectively. 

The  dendrogram  in  Figure  9 differs  from  that  in  Figure  8 
in  several  respects.  The  former  dendrogram  contains  fewer 
unique  synapomorphies  (three  rather  than  six)  and  more  par- 
allelisms (ten  rather  than  five),  but  it  has  no  reversals.  A five- 
way division  occurs  above  the  first  synapomorphy.  The  three 
characters  involved  in  unique  synapomorphies  (1,7,  11)  fall 


16  Contrikiutions  in  Science,  Number  345 


Wake  and  Elias:  Tropical  Salamanders 


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supergenus 

Bolitoglossa 

Figure  9.  An  alternative  tree  of  the  genera  in  the  tribe  Bolitoglos- 
sini.  There  are  no  reversals  in  this  tree,  but  all  parallelisms  (asterisks) 
are  indicated. 


into  Hecht  and  Edwards’s  (1976a)  third  and  fourth  cate- 
gories, the  kinds  of  characters  that  they  consider  to  have 
relatively  high  systematic  value.  The  stem  uniting  Lineatri- 
ton,  Parvimolge,  and  Thorius  has  three  synapomorphies.  For 
illustrative  purposes,  we  have  indicated  all  parallelisms  in 
Figure  9,  and  it  can  be  seen  that  two  of  these  three  synapo- 
morphies evolved  in  parallel  with  other  stems.  The  stem 
uniting  Bradytriton,  Nototriton,  Oedipina,  and  Bolitoglossa 
also  has  three  synapomorphies  (4,  6,  9),  but  none  of  them  is 
unique. 

The  tropical  plethodontid  genera  constitute  a highly  de- 
rived lineage.  The  species  are  so  specialized,  in  fact,  that  little 
further  morphological  innovation  seems  possible.  Rather, 
variation  on  similar  themes  has  occurred  repeatedly,  with 
the  result  that  characters  useful  in  the  taxonomy  of  other 
salamander  groups  have  been  subject  to  extensive  parallel 
evolution.  We  have  long  known  that  such  features  as  elon- 
gation of  body  and  tail,  diminution  in  body  size,  changes  in 


relative  limb  length  and  foot  size,  and  increased  digital  web- 
bing have  evolved  several  times  in  the  group  (Wake,  1966; 
Wake  and  Fynch,  1976;  Alberch  and  Alberch,  1981).  Ap- 
parently the  evolution  of  other  morphological  traits  is  also 
complex,  and  close  evolutionary  interlinking  of  seemingly 
unrelated  traits  is  a possibility  (see  Alberch  et  al.,  1979;  Al- 
berch and  Alberch,  1981).  The  incorporation  of  biochemical 
and  immunological  approaches  to  phyletic  analysis  holds 
promise  for  unraveling  the  complex  history  of  this  group, 
but  the  long  evolutionary  history  of  the  neotropical  sala- 
manders poses  serious  problems  for  all  known  analytical 
techniques.  At  one  time,  the  tropical  salamanders  were  con- 
sidered to  form  a reasonably  compact  congeneric  group  ( Dunn, 
1926).  The  intervening  years  have  seen  a steady  erosion  of 
this  concept  as  new  salamanders  have  been  discovered  and 
novel  analytical  techniques  have  been  applied.  Based  pri- 
marily on  its  possession  of  a common  tongue  projection 
mechanism  (Lombard  and  Wake,  1 977),  the  supergenus  Boli- 
toglossa does  appear  to  be  monophyletic,  but  the  group  is 
more  ancient  and  highly  diverse  than  was  thought  previ- 
ously. The  present  effort  is  but  one  more  developmental  stage 
in  our  attempt  to  understand  the  complex  history  of  the 
tropical  salamanders. 

ACKNOWLEDGMENTS 

Work  reported  in  this  paper  has  been  under  way  for  several 
years  and  has  involved  examination  of  specimens  housed  in 
a number  of  collections.  We  especially  thank  the  curators  of 
the  collections  of  the  Museum  of  Natural  History,  University 
of  Kansas;  National  Museum  of  Natural  History;  American 
Museum  of  Natural  History;  Field  Museum  of  Natural  His- 
tory; Museum  of  Comparative  Zoology,  Harvard  University; 
Museum  of  Zoology,  University  of  Michigan;  and  the  Nat- 
ural History  Museum  of  Los  Angeles  County  for  loan  of 
specimens  and  for  hospitality  and  access  to  their  collections 
during  our  visits.  Monica  Frelow  assisted  in  several  phases 
of  our  study.  Gene  M.  Christman  of  the  Museum  of  Verte- 
brate Zoology  and  James  Hendel  of  the  Scientific  Photo- 
graphic Laboratory,  University  of  California,  Berkeley,  as- 
sisted in  preparation  of  the  figures.  We  have  benefitted  from 
discussions  concerning  several  aspects  of  this  study  with  Ar- 
den H.  Brame,  James  Kezer,  James  F.  Lynch,  Stanley  Ses- 
sions, Marvalee  H.  Wake,  and  Kay  P.  Yanev.  We  are  in- 
debted to  several  colleagues  for  extensive  help  in  our  field 
work,  especially  Jeremy  L.  Jackson,  James  F.  Lynch,  Theo- 
dore J.  Papenfuss,  Robert  Seib,  and  H.  Bradley  Shaffer,  but 
also  to  others  too  numerous  to  mention.  The  work  of  this 
laboratory  on  tropical  salamanders  has  been  supported  by 
the  National  Science  Foundation  (current  grant  DEB-78 
03008).  Finally,  we  thank  the  Guatemalan  Forest  Service  for 
making  it  possible  for  us  to  work  in  Guatemala,  and  the 
Direccion  General  de  la  Fauna  Silvestre  for  collecting  permits 
for  Mexico. 


LITERATURE  CITED 

Alberch,  P.,  and  J.  Alberch.  1981.  Heterochronic  mecha- 
nisms of  morphological  diversification  and  evolutionary 


Contributions  in  Science,  Number  345 


Wake  and  Elias:  Tropical  Salamanders  17 


change  in  the  neotropical  salamander  Bo/itoglossa  oc- 
cidentalis  (Amphibia:  Plethodontidae).  J.  Morph.  167: 
249-264. 

Alberch,  P„  S.J.  Gould,  G.F.  Oster,  and  D.B.  Wake.  1979. 
Size  and  shape  in  ontogeny  and  phylogeny.  Paleobiology 
5:296-317. 

Arnold,  S.J.  1977.  The  evolution  of  courtship  behavior  in 
new  world  salamanders  with  some  comments  on  old 
world  salamandrids.  Pages  141-183  in  D.H.  Taylor  and 
S.E.  Guttman  (eds.).  The  reproductive  biology  of  am- 
phibians, Plenum  Press,  New  York  City. 

Baird,  I.L.  195  1 . An  anatomical  study  of  certain  salaman- 
ders of  the  genus  Pseudoeurycea.  Univ.  Kansas  Sci.  Bull 
34,  pt.  1:221-265. 

Brame,  A.H.,  Jr.  1968.  Systematics  and  evolution  of  the 
Mesoamerican  salamander  genus  Oedipina.  J.  Herpet. 
2:1-64. 

Cope,  E.D.  1869.  A review  of  the  species  of  the  Pletho- 
dontidae and  Desmognathidae.  Proc.  Acad.  Nat.  Sci. 
Philadelphia  1869:93-118. 

Dunn,  E.R.  ,1926.  The  salamanders  of  the  family  Pletho- 
dontidae. Smith  College,  Northampton,  Mass.,  viii  + 
441  pp. 

Edwards,  J.L.  1976.  Spinal  nerves  and  their  bearing  on 
salamander  phylogeny.  J.  Morph.  148:305-328. 

Eldredge,  N.,  and  J.  Cracraft.  1980.  Phylogenetic  patterns 
and  the  evolutionary  process.  Columbia  Univ.  Press,  New 
York  City,  viii  + 349  pp. 

Elias,  P.  Salamanders  of  the  northwestern  highlands  of  Gua- 
temala. Contrib.  Sci.  Nat.  Hist.  Mus.  Los  Angeles  Co., 
in  press. 

Elias,  P.,  and  D.  B.  Wake.  1983.  Nyctanolis  pernix,  a new 
genus  and  species  of  plethodontid  salamander  from 
northwestern  Guatemala  and  Chiapas,  Mexico.  Pages 
1-12  in  A.G.J.  Rhodin  and  K.  Miyata  (eds.).  Advances 
in  herpetology  and  evolutionary  biology’:  Essays  in  honor 
of  Ernest  E.  Williams,  The  Museum  of  Comparative 
Zoology,  Cambridge,  Mass. 

Farris,  J.S.  1970.  Methods  for  computing  Wagner  trees. 
Systematic  Zoology  19:83-92. 

Freeman,  H.L.  1977.  Systematics  and  evolutionary  trends 
of  the  Mexican  salamander  genus  Thorius  Cope,  1869 
(Amphibia,  Plethodontidae).  Ph.D.  thesis,  Rutgers  Univ. 

Hanken.J.  1980.  Morphological  and  genetic  investigations 
of  miniaturization  in  salamanders  (genus  Thorius).  Ph.D. 
dissertation,  Univ.  California,  Berkeley. 

Hecht,  M.K.  1976.  Phylogenetic  inference  and  methodol- 
ogy as  applied  to  the  vertebrate  record.  Vol.  9,  pp.  335— 
363,  in  M.K.  Hecht,  W.C.  Steere,  and  B.  Wallace  (eds.). 
Evolutionary  biology.  Plenum  Press,  New  York  City. 

Hecht,  M.K.,  and  J.L.  Edwards.  1976a.  The  determination 
of  parallel  or  monophyletic  relationships:  The  proteid 
salamanders— a test  case.  Am.  Nat.  1 10:653-677. 

. 1 976b.  The  methodology  of  phylogenetic  inference 

above  the  species  level.  Pages  3-5  1 in  M.K.  Hecht,  P.C. 
Goody,  and  B.M.  Hecht  (eds.).  Major  patterns  in  ver- 
tebrate evolution,  Plenum  Press,  New  York  City. 

Larson,  A.,  D.B.  Wake,  L.R.  Maxson,  and  R.  Highton.  1981. 


A molecular  phylogenetic  perspective  on  the  origins  of 
morphological  novelties  in  the  salamanders  of  the  Tribe 
Plethodontini  (Amphibia,  Plethodontidae).  Evolution 
35:405-422. 

Leon,  P.C.,  and  J.  Kezer.  1978.  Localization  of  5S  RNA 
genes  on  chromosomes  of  plethodontid  salamanders. 
Chromosoma  65:213-230. 

Lombard,  R.E.,  and  D.B.  Wake.  1977.  Tongue  evolution 
in  the  lungless  salamanders,  family  Plethodontidae.  II. 
Function  and  evolutionary  diversity.  J.  Morph.  1 53:39— 
80. 

Lynch,  J.F.,  and  D.B.  Wake.  1975.  Systematics  of  the  Chi- 
ropterotriton  bromeliacia  group  (Amphibia:  Caudata), 
with  description  of  two  new  species  from  Guatemala. 
Contrib.  Sci.  Nat.  Hist.  Mus.  Los  Angeles  Co.  265:1- 
45. 

. 1978.  A new  species  of  Chiropterotriton  (Amphib- 
ia: Caudata)  from  Baja  Verapaz,  Guatemala,  with  com- 
ments on  relationships  among  Central  American  mem- 
bers of  the  genus.  Contrib.  Sci.  Nat.  Hist.  Mus.  Los 
Angeles  Co.  294:1-22. 

Marlow,  R.W.,  J.M.  Brode,  and  D.B.  Wake.  1979.  A new 
salamander,  genus  Batrachoseps,  from  the  Inyo  Moun- 
tains of  California,  with  a discussion  of  relationships  in 
the  genus.  Contrib.  Sci.  Nat.  Hist.  Mus.  Los  Angeles  Co. 
308:1-17. 

Maxson,  L.R.,  and  D.B.  Wake.  1981.  Albumin  evolution 
and  its  phylogenetic  implications  in  the  plethodontid 
salamander  genera  Pseudoeurycea  and  Chiropterotriton. 
Herpetologica  37:109-1 17. 

Rabb.  G.B.  1955.  A new  salamander  of  the  genus  Parvi- 
mo/ge  from  Mexico.  Breviora,  Mus.  Comp.  Zool,  42: 
1-9. 

. 1956.  A new  plethodontid  salamander  from  Nuevo 

Leon,  Mexico.  Fieldiana-Zoology,  Chicago  39:1  1-20. 

. 1958.  On  certain  Mexican  salamanders  of  the  pleth- 
odontid genus  Chiropterotriton.  Occ.  Pap.  Mus.  Zool. 
Univ.  Michigan  587:1-37. 

. 1965.  A new  salamander  of  the  genus  Chiroptero- 
triton (Caudata:  Plethodontidae)  from  Mexico.  Breviora, 
Mus.  Comp.  Zool.,  235:1-8. 

Robinson,  D.C.  1976.  Anewdwarfsalamanderofthegenus 
Bolitoglossa  (Plethodontidae)  from  Costa  Rica.  Proc. 
Biol.  Soc.  Wash.  89:289-294. 

Smith,  H.M.,  and  R.B.  Smith.  1976.  Synopsis  of  the  her- 
petofauna  of  Mexico,  vol.  IV:  Source  analysis  and  index 
for  Mexican  amphibians.  John  Johnson,  North  Ben- 
nington, Vermont. 

Tanner,  W.W.  1950.  A new  genus  of  plethodontid  sala- 
mander from  Mexico.  Great  Basin  Natur.  10:37-44. 

. 1952.  A comparative  study  of  the  throat  muscu- 
lature of  the  Plethodontidae  of  Mexico  and  Central 
America.  Univ.  Kans.  Sci.  Bull.  34,  pt.  2:583-677. 

Taylor,  E.H.  1940.  New  salamanders  from  Mexico,  with  a 
discussion  of  certain  known  forms.  Univ.  Kans.  Sci. 
Bull.  26:407-439. 

. 1941.  New  plethodont  salamanders  from  Mexico. 

Herpetologica  2:57-65. 


18  Contributions  in  Science,  Number  345 


Wake  and  Elias:  Tropical  Salamanders 


. 1944.  The  genera  of  plethodont  salamanders  in 

Mexico,  pt.  1.  Univ.  Kans.  Sci.  Bull.  30:189-232. 

Trufelli,  G.T.  1 954.  A macroscopic  and  microscopic  study 
of  the  mental  hedonic  gland-clusters  of  some  pletho- 
dontid  salamanders.  Univ.  Kans.  Sci.  Bull.  36,  pt.  1:3- 
39. 

Wake,  D.B.  1966.  Comparative  osteology  and  evolution  of 
the  lungless  salamanders,  family  Plethodontidae.  Mem- 
oirs, So.  Calif.  Acad.  Sci.  4:1-1  1 1. 

. 1970.  Aspects  of  vertebral  evolution  in  the  modem 

Amphibia.  Forma  et  Functio  3:33-60. 

. 1 972.  Emmett  Reid  Dunn  and  the  development  of 

scientific  studies  of  salamander  biology.  Pages  iii-x  in 
Introduction  to  reprint  of  E.R.  Dunn,  Salamanders  of 
the  family  Plethodontidae,  facsimile  reprints.  Society  for 
the  Study  of  Amphibians  and  Reptiles,  Athens,  Ohio. 


Wake,  D.B.,  and  A.H.  Brame,  Jr.  1963.  The  status  of  the 
plethodontid  salamander  genera  Bolitoglossa  and  Mag- 
nadigita.  Copeia  1963:382-387. 

. 1969.  Systematics  and  evolution  of  neotropical  sal- 
amanders of  the  Bolitoglossa  helmrichi  group.  Contrib. 
Sci.  Nat.  Hist.  Mus.  Los  AngelesGo.  175:1  — 40. 

Wake,  D.B.,  and  I.G.  Dresner.  1967.  Functional  mor- 
phology and  evolution  of  tail  autonomy  in  salamanders. 
J.  Morph.  122:265-306. 

Wake,  D.B.,  and  J.F.  Lynch.  1976.  The  distribution,  ecol- 
ogy, and  evolutionary  history  of  plethodontid  salaman- 
ders in  tropical  America.  Sci.  Bull.  Nat.  Hist.  Mus.  Los 
Angeles  Co.  25:1-65. 


Submitted  6 October  1981;  accepted  15  December  1982. 


Contributions  in  Science,  Number  345 


Wake  and  Elias:  Tropical  Salamanders  19 


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mm 


Number  346 

7 October  I9S3 


, , . 


THE  CRANIAL  MORPHOLOGY  ilF  THE  FOSSIL  BEAVER 
DIPOIDES  SMITH. I (ROBENITA:  MAMMALIA) 


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Craig  C.  Black,  Museum  Director 
Donald  Chapul 

Daniel  M.  Cohen,  Committee  Chairman 
John  M.  Harris 
Charles  L.  Hogue 

Robin  A.  Simpson,  Managing  Editor 
Gary  D.  Wallace 
Edward  C.  Wilson 


Printed  at  Allen  Press,  Inc.,  Lawrence.  Kansas 


THE  CRANIAL  MORPHOLOGY  OF  THE  FOSSIL  BEAVER 
DIPOIDES  SMITHI  (RODENTIA:  MAMMALIA) 

Hugh  M.  Wagner1 


ABSTRACT.  The  cranial  morphology  of  the  genus  of  fossil  beaver, 
Dipoides,  has  never  been  described  or  compared  with  that  of  other 
fossil  and  extant  beavers.  An  extremely  well-preserved  skull  re- 
covered from  north-central  Oregon  in  1974  allows  for  identification 
of  nearly  all  features  of  the  cranium.  Variation  in  skull  morphology 
in  rodents  is  generally  recognizable  at  the  generic  level,  and  intra- 
specific variation  is  usually  reflected  in  differences  in  the  dentition. 
The  skull  of  Dipoides  closely  resembles  that  of  Eucastor  and  Cas- 
toroides  and  differs  noticeably  from  that  of  Castor.  The  regions  of 
the  skull  with  similarities  to  Dipoides,  Eucastor,  and  Castoroides  are 
( 1 ) the  bones  of  the  lacrimal  region;  (2)  the  arrangement  of  foramina 
in  the  basicranium  and  interorbital  region;  and  (3)  the  development 
of  the  alisphenoid  wing  of  the  internal  pterygoid  fossa.  Castor  differs 
in  these  three  areas.  Close  examination  of  the  crania  of  these  four 
genera  confirms  that  Eucastor,  Dipoides,  and  Castoroides  represent 
one  group  of  beavers  that  has  been  phyletically  distinct  from  that 
leading  to  the  extant  beaver.  Castor,  since  the  early  Miocene. 

INTRODUCTION 

Species  in  the  genus  Dipoides  Jager,  1835,  are  small  beavers 
that  lived  during  the  Hemphillian  Mammal  Age  in  North 
America.  The  genus  appears  to  have  evolved  from  Eucastor 
Leidy,  1 858,  during  the  late  Miocene,  approximately  8.5  mya 
(Wagner,  1981). 

The  genus  Dipoides  was  first  described  by  G.F.  Jager  in 
1835  from  late  Miocene  deposits  of  Germany.  Subsequently, 
other  species  of  Dipoides  have  been  described  from  late  Mio- 
cene deposits  of  Eurasia  and  North  America.  R.A.  Stirton 
(1935)  reviewed  the  Tertiary  beavers,  presenting  a phylogeny 
based  on  dental  characters,  with  Dipoides  in  the  lineage  lead- 
ing to  Castoroides  Foster,  1838.  Later,  Olson  (1940)  under- 
took a comparative  study  of  the  cranial  foramina  of  the  Cas- 
toridae.  He  described  and  compared  the  cranial  foramina  of 
Paleocastor  Leidy,  1869,  Eucastor,  Castoroides,  and  Castor 
Linneaus,  1758,  noting  evolutionary  trends  in  foramina  with- 
in the  Castoridae.  Dipoides  was  not  included  in  that  study. 

J.A.  Shotwell  (1955)  reviewed  the  genus  Dipoides,  basing 
his  taxonomic  conclusions  on  the  dental  morphology,  pri- 


1. Department  of  Physics,  University  of  California,  Irvine,  Cal- 
ifornia 92717,  and  Research  Associate,  Vertebrate  Paleontology, 
Natural  History  Museum  of  Los  Angeles  County. 


marily  P4.  He  concluded  with  the  description  of  a new  species, 
D.  smithi  Shotwell,  1955,  from  McKay  Reservoir  in  north- 
central  Oregon.  Stirton  (1965)  described  the  cranial  mor- 
phology of  Castoroides  ohioensis  Foster,  1838,  in  detail,  in- 
creasing the  knowledge  of  this  giant  castorid.  The  specimen 
of  D.  smithi  that  I describe  in  the  present  paper  was  recovered 
from  the  type  locality  of  the  species  at  McKay  Reservoir  (Fig. 
1 ) in  1974  and  provides  new  evidence  concerning  the  cranial 
morphology  of  the  genus  and  helps  clarify  its  relationship  to 
other  genera  of  castorids. 

METHODS  AND  MATERIALS 

The  abbreviations  used  in  the  text  and  tables  are  CNHM 
(Chicago  Natural  History  Museum),  UO  (University  of  Or- 
egon), and  UCMP  (University  of  California  Museum  of  Pa- 
leontology). All  measurements  are  in  millimeters.  The  com- 
parative specimens  are: 

Eucastor  cf.  E.  tortus  Leidy,  1858,  CNHM  P15787,  Big 
Springs  Canyon  local  fauna,  Bennett  County,  South  Da- 
kota, Ogallala  Group,  Clarendonian. 

Castoroides  ohioensis  Foster,  1838,  CNHM  P3800,  Gravel 
Pit  near  Bellflower,  McClean  County,  Illinois  Pleisto- 
cene. 

Castor  canadensis  Kuhl,  1820,  UCMP  126-158,  South 
Dakota,  Recent. 

Dipoides  smithi  Shotwell,  1955,  UCMP  113422,  McKay 
Reservoir  local  fauna,  Umatilla  County,  Oregon,  ? Shu- 
tler  Formation,  Hemphillian  (Figs.  2,  3 and  4). 

SYSTEMATICS 

Family  Castoridae  Gray,  1 92 1 
Subfamily  Castoroidinae  Trouessant,  1888 
Genus  Dipoides  Jager,  1835 

Dipoides  smithi  Shotwell,  1955 

Figures  2,  3,  and  4 

REFERRED  MATERIAL.  UCMP  113422,  nearly  com- 
plete skull  lacking  only  the  projection  of  the  premaxillaries 


Contributions  in  Science,  Number  346,  pp.  1-6 
Natural  History  Museum  of  Los  Angeles  County,  1983 


ISSN  0459-8113 


Figure  1.  The  skull  of  Dipoides  smithi  was  recovered  from  UCMP 
locality  V74163  at  McKay  Reservoir  in  Umatilla  County  in  north- 
central  Oregon,  designated  by  an  X. 


and  incisors  (Figs.  2 and  3);  UCMP  113433,  left  dentary 
fragment  with  P4,  M,_2  (Fig.  4),  both  from  locality  UCMP 
V74 1 63  = U02222,  McKay  Reservoir,  Umatilla  County, 
Oregon,  ? Shutler  Formation,  Hemphillian. 

DESCRIPTION  AND  COMPARISON.  The  dental 
morphology  of  UCMP  1 13422  agrees  with  the  diagnosis  of 
D.  smithi  (Shotwell,  1955).  Upper  molars  and  premolars  lack 
parastria  and  have  S-shaped  occlusal  patterns  characteristic 
of  this  species.  In  the  sample  of  P4’s  of  D.  smithi  from  McKay 
Reservoir,  16  percent  of  24  P4’s  had  complete  or  vestigial 
parastriids  and  paraflexids. 

The  incisors  and  most  of  the  ventral  region  of  the  pre- 
maxillary bone  are  absent  in  this  specimen.  The  dorsal  sur- 
face of  the  skull  is  long  and  relatively  flat.  Over  the  orbits 
the  frontal  bone  forms  a broad,  Hat  surface  (Fig.  2).  The  nasal 
region  is  straight,  with  neither  the  lateral  nor  dorsal  inflation 
of  Castor  canadensis.  A small  protuberance  exists  on  the 
dorsal  lip  of  the  fossa  for  the  masseter  profundis  anterior 
muscle  at  the  premaxillary-maxillary  suture.  This  protuber- 
ance is  similar  to  that  in  Castoroides  ohioensis ; Eucastor  has 
only  a slight  one  and  Castor  lacks  one.  Frontoparietal  crests 
arise  above  the  orbits  and  extend  posteriorly  along  the  fron- 
tal-parietal suture,  where  they  join  to  form  a strong  median 
sagittal  crest  between  the  parietal  bones.  The  lambdoidal 
crests  are  well  developed,  extending  laterally  nearly  to  the 
tip  of  the  mastoid  processes. 

The  anterior  and  posterior  temporal  foramina  are  both 
present  on  the  parietal-squamosal  suture  and  in  the  same 
position  as  in  Eucastor  and  Castoroides.  Approximately  ninety 
percent  of  the  surfaces  of  the  parietal  bones  are  covered  with 
rugosities.  Eucastor  and  Castoroides  also  have  similar  ru- 
gosities although  not  as  extensive  anteriorly  as  in  this  spec- 
imen of  Dipoides.  In  Castor,  such  rugosities  are  not  as  well 
developed.  The  dorsal  surface  of  the  cranium  in  Dipoides  is 
more  inflated  than  in  Eucastor  but  not  as  inflated  as  in  Cas- 
tor. The  occipital  plate  intersects  a plane  extended  posterior 


Figure  2.  Dipoides  smithi : Shotwell,  1955,  UCMP  1 1 3422,  McKay 
Reservoir  local  fauna,  UCMP  locality  V74 1 63,  nearly  complete  skull 
lacking  the  incisor  region;  A,  dorsal  view;  B,  right  lateral  view;  C, 
ventral  view;  scale  indicated. 

from  the  palate  at  approximately  seventy  degrees.  By  com- 
parison, in  Eucastor,  Castoroides,  and  Castor,  the  occipital 
plate  intersects  the  same  plane  at  nearly  ninety  degrees.  A 
small  mastoid  foramen  pierces  the  mastoid  bone  slightly  lat- 
eral to  the  mastoid-supraoccipital  suture.  In  size  and  position 
the  mastoid  foramen  is  very  similar  to  that  in  Castoroides. 
In  Castor,  this  foramen  is  much  larger  and  situated  on  the 
mastoid-supraoccipital  suture.  The  occipital  condyles  appear 
very  similar  to  those  in  Castor  and  do  not  show  the  transverse 
flattening  of  those  of  Castoroides. 


2 Contributions  in  Science,  Number  346 


Wagner:  Cranial  Morphology  of  Dipoides  smithi 


Anterior  and  ventral  to  the  infraorbital  foramen  there  is  a 
very  prominent  masseter  superficialis  process  (Fig.  3,  B).  A 
prominent  flange  or  sheath  of  bone  extends  dorsally  from  the 
lateral  edge  of  this  process,  closing  off  the  ventral  region  of 
the  infraorbital  foramen  from  the  masseter  lateralis  profun- 
dus anterior  fossa  (Fig.  3,  B).  This  flange  bends  posteriorly 
toward  its  dorsal  limit  and  does  not  extend  above  the  in- 
fraorbital foramen.  The  morphology  of  the  masseteric  su- 
perficialis process  and  infraorbital  complex  in  Dipoides  is 
nearly  identical  to  that  of  Castoroides.  In  Eucastor  the  mas- 
seteric superficialis  process  is  not  well  developed,  and  the 
structure  of  the  infraorbital  region  resembles  that  of  Castor. 

The  zygomatic  arch  is  wide,  but  not  to  the  extreme  seen 
in  Castor  or  Castoroides.  The  orbit  of  Dipoides  is  propor- 
tionally larger  than  in  Castor  and  Castoroides  and  appears 
to  be  directed  more  laterally  than  in  Castor.  The  structure 
of  the  zygomatic  arch  and  lacrimal  region  is  similar  to  that 
in  Castoroides  and  Eucastor  and  differs  from  that  in  Castor. 
The  contribution  that  the  jugal  bone  makes  to  the  zygomatic 
arch  of  Dipoides,  Eucastor,  and  Castoroidesd s less  than  in 
Castor.  In  the  former  three  genera,  the  jugal-maxilla  suture 
is  on  the  lateral  surface  of  the  zygomatic  arch  as  contrasted 
to  Castor  where  this  suture  is  in  the  lacrimal  area.  The  an- 
terior margin  of  the  orbit  is  composed  of  the  maxilla,  because 
the  jugal  does  not  extend  into  the  lacrimal  region.  In  Castor, 
however,  the  jugal-maxilla  suture  extends  anteriorly  on  the 
dorsal  surface  of  the  zygomatic  arch,  and  the  jugal  meets  the 
lacrimal  in  the  anterior  region  of  the  orbit. 

Dipoides  has  a simple  lacrimal  region  in  which  only  the 
frontal  and  maxilla  meet  on  the  dorsal  surface  of  the  skull. 
The  lacrimal  bone  does  not  have  a dorsal  component  in  this 
specimen.  The  structure  of  the  lacrimal  region  in  Dipoides 
and  Castoroides  is  similar  by  having  only  the  two  dorsal 
elements.  Eucastor  differs  slightly  from  Dipoides  and  Cas- 
toroides, because  a small  portion  of  the  lacrimal  bone  is 
visible  on  the  dorsal  surface  at  the  maxillary-frontal  suture. 
Castor  has  a much  more  complicated  dorsal  lacrimal  area  in 
which  the  jugal,  maxillary,  lacrimal,  and  frontal  bones  all 
meet. 

The  pattern  of  foramina  within  the  inner  orbit  of  Dipoides 
is  very  similar  to  that  in  Eucastor  and  distinctly  different 
from  the  pattern  in  either  Castoroides  or  Castor.  However, 
the  resemblance  to  Castoroides  is  greater  than  to  Castor.  The 
sphenoid  foramen  is  the  anterior  foramen,  piercing  the  max- 
illary in  an  anterior  direction.  Directly  behind  the  sphenoid 
foramen  is  a slightly  smaller  one  that  enters  the  cranium 
medially.  This  foramen  completely  penetrates  the  cranium 
as  does  the  presphenoid  canal  of  Olson  (1940)  (alisphenoid- 
rotundum  foramen  of  Stirton  (1965)).  This  foramen  appears 
to  be  an  anterior  presphenoid  canal.  Directly  above  the  an- 
terior presphenoid  canal  on  the  orbitosphenoid-frontal  su- 
ture is  the  ethmoid  foramen.  Posterior  and  ventral  to  the 
ethmoid  foramen  is  a well-developed  optic  foramen  entirely 
enclosed  in  the  orbitosphenoid  bone.  A large  posterior  pre- 
sphenoid canal  is  situated  posterior  to  the  optic  foramen  in 
a deep  sulcus  at  the  anterior  end  of  the  sphenoidal  fissure. 
Near  the  anteroposterior  margin  of  the  alisphenoid  wing  of 
the  internal  pterygoid  fossa  is  the  masticatory-buccinator 


Figure  3.  Dipoides  smithi,  Shotwell,  1955,  skull  with  structures 
labeled,  UCMP  1 13422;  A,  dorsal  view;  B,  right  lateral  view;  C, 
ventral  view;  scale  indicated  roman  type  represents  bones  and  struc- 
ture, italics  are  foramina.  Abbreviations:  af,  alisphenoid  foramen; 
als,  alisphenoid;  apf,  anterior  palatine  foramen;  atf  anterior  tem- 
poral foramen;  awipf,  alisphenoid  wing  of  internal  pterygoid  process; 
bo,  basioccipital;  earn,  external  auditory  meatus;  ef.  ethmoid  fora- 
men; eo,  exoccipital;  fr,  frontal;  hf,  hypoglossal  foramen;  iof,  in- 
fraorbital foramen;  ip,  interparietal;  j,  jugal;  m,  mastoid;  mf,  mastoid 
foramen;  mlf-fo,  median  lacerate  foramen-foramen  ovale;  mp,  mas- 
toid process;  ms-bcf  masticatory-buccinator  foramen;  msp,  mas- 
seteric superficialis  process;  mx,  maxillary;  n,  nasal;  of,  optic  fora- 
men; p,  parietal;  pgf,  postglenoid  foramen;  pi,  palatine;  pp,  paroccipital 
process;  prsp.  can.,  presphenoid  canal;  ps,  premaxillary;  pt,  ptery- 
goid; ptf,  posterior  temporal  foramen;  sf,  stylomastoid  foramen;  so, 
supraoccipital;  sphf  sphenoidal  foramen;  sq,  squamosal;  ty,  tym- 
panic; vc,  ventral  choana. 


Contributions  in  Science,  Number  346 


Wagner:  Cranial  Morphology  of  Dipoides  smithi  3 


I cm. 


Figure  4.  Dipoides  smithi,  UCMP  1 13433,  UCMP  V74163,  col- 
lected near  the  skull  figured  in  this  paper.  Left  dentary  fragment  with 
P4,M,_2  showing  normal  occlusal  wear.  This  specimen  has  the  more 
typical  P4  of  D.  smithi,  where  no  paraflexid  is  present. 

foramen.  Below  the  masticatory-buccinator  foramen,  a por- 
tion of  the  alisphenoid  canal  is  visible,  as  it  passes  through 
the  alisphenoid  bone.  Anteroventral  to  the  alisphenoid  canal 
is  a large  foramen  that  appears  to  enter  the  pulp  cavity  of 
M3.  The  pattern  of  foramina  anterior  to  and  including  both 
presphenoid  canals  in  Dipoides  is  very  similar  to  that  in 
Eucasior  and  Castor.  Eucastor  resembles  Dipoides  by  having 


the  multiple  presphenoid  canals.  By  contrast,  in  Castoroides, 
the  presphenoid  canal  is  single  and  proportionally  much 
smaller  and  more  posterior  in  its  position. 

Posterior  to  the  presphenoid  canal,  the  morphology  of  the 
alisphenoid  in  Dipoides  does  not  resemble  that  in  Castor  but 
is  very  similar  to  that  in  Castoroides.  In  Dipoides  and  Cas- 
toroides, the  alisphenoid  forms  a smooth  lateral  surface  of 
the  sphenoid  fissure  from  the  anterior  sulcus  of  the  presphe- 
noid canal  posterior  to  the  masticatory-buccinator  foramen. 
The  masticatory-buccinator  foramen  is  directed  posteriorly 
through  the  margin  of  the  alisphenoid.  In  Castor,  part  of  the 
alisphenoid  forms  a strap-like  structure,  forming  the  lateral 
surface  of  the  alisphenoid  canal.  The  masticatory-buccinator 
foramen  and  alisphenoid  foramen  share  a common  orifice 
that  is  at  the  posterior  end  of  the  straplike  structure  of  the 
alisphenoid.  This  common  foramen  is  directed  anteriorly, 
entering  directly  into  the  sphenoid  fissure.  The  alisphenoid 
foramen  pierces  the  broad  alisphenoid  wing  of  Dipoides  and 
Castoroides,  and  a branch  of  the  trigeminal  nerve  (V2)  enters 
the  skull  through  a bone  canal  enclosed  in  the  alisphenoid 
wing  of  the  internal  pterygoid.  Castor  differs  in  that  the  ali- 
sphenoid foramen  does  not  pass  through  the  alisphenoid  wing, 
but  the  trigeminal  nerve  enters  via  the  alisphenoid  foramen. 
This  region  of  the  skull  of  the  specimen  of  Eucastor  was  not 
well  enough  preserved  to  allow  comparison. 

The  postglenoid  foramen  is  large  and  nearly  equal  in  size 


Table  1.  A comparison  of  cranial  characters  in  Eucastor  cf.  E.  tortus,  CNHM  PI 5787;  Dipoides  smithi,  UCMP  113422;  Castoroides  ohioensis, 
CNHM  P3800;  and  Castor  canadensis,  UCMP  126-158. 


Characters 

E.  cf.  E.  tortus 

D.  smithi 

C.  ohioensis 

C.  canadensis 

Protuberance  on  dorsal  lip  of  anterior 

slight 

slight 

well-developed 

absent 

masseteric  fossa 

Temporal  foramina  present 

yes 

yes 

yes 

yes 

Rugosities  present  on  parietal  bone 

slight 

well-developed 

well-developed 

slight 

Inflation  of  cranium 

slight 

slight 

slight 

well-developed 

Mastoid  foramen 

small 

small 

small 

large 

Masseteric  superficialis  process  well  developed 

no 

yes 

yes 

yes 

Masseteric  superficialis  process  anterior  to 

no 

yes 

yes 

no 

anterior  masseteric  fossa 

Lateral  vertical  flange  on  infraorbital  foramen 

yes 

no 

no 

yes 

Parallel  tooth  row  on  maxillary 

yes 

no 

no 

yes 

Dorsal  choana  or  invagination  on  pterygoid  or 

no 

no 

yes 

yes 

basioccipital  bones 

Posterior  carotid  foramen 

absent 

absent 

present 

present 

Alisphenoid  foramen  pierces  medial 

not  known 

yes 

yes 

no 

alisphenoid  wing  of  internal  pterygoid  fossa 

Jugal  extends  into  lacrimal  region 

no 

no 

no 

yes 

Multiple  presphenoid  canals 

yes 

yes 

no 

yes 

Bulla  highly  inflated 

yes 

yes 

no 

yes 

Mastoid  process  extends  beyond  external 

no 

no 

no 

yes 

auditory  meatus 

4 Contributions  in  Science,  Number  346 


Wagner:  Cranial  Morphology  of  Dipoides  smithi 


to  the  external  auditory  meatus.  The  external  auditory  me- 
atus is  posterior  and  ventral  to  the  postglenoid  foramen  and 
is  appressed  posteriorly  against  the  mastoid  process  and  di- 
rected laterally.  Its  anterior  lip  has  the  greatest  lateral  exten- 
sion. The  condition  of  the  postglenoid  foramen  and  external 
auditory  meatus  is  very  similar  to  that  of  Castoroides.  Castor 
and  Eucastor  differ,  having  a much  smaller  postglenoid  fo- 
ramen; the  meatus  has  a more  dorsolateral  orientation,  and 
its  ventral  lip  has  the  greatest  lateral  extension.  In  Dipoides 
and  Castor,  the  meatus  projects  laterally  beyond  the  mastoid 
process.  In  Castoroides,  it  does  not. 

The  internal  pterygoid  fossa  is  broad,  bounded  laterally  by 
the  alisphenoid  wing  and  medially  by  the  palatal  wing  of  the 
pterygoid,  which  is  a septum  supporting  the  hamular  process. 
The  alisphenoid  foramen  is  present  just  anterior  to  the  con- 
joined median  lacerate  foramen  and  foramen  ovale  on  the 
alisphenoid  wing  of  the  internal  pterygoid  fossa  (Fig.  3,  C). 
The  surface  of  the  pterygoid  fossa  has  depressions.  The  ven- 
tral choana  is  triangular  in  outline,  bounded  laterally  by  the 
palatal  wings  of  the  pterygoid.  In  Castoroides  the  outline  of 
the  ventral  choana  is  similar  to  that  of  Dipoides,  but  in  Castor 
it  is  nearly  circular. 

There  is  a rugosity  on  the  anterior  median  border  of  the 
bulla,  resembling  that  of  Castor,  where  the  hamuiar  process 
meets  the  tympanic.  In  Castor  these  rugosities  extend  farther 
posteriorly,  so  it  seems  probable  that  the  hamular  processes 
of  Dipoides  were  not  as  long  as  in  Castor.  The  overall  struc- 
ture of  the  internal  pterygoid  fossa  in  Dipoides  resembles  that 
in  Castoroides,  in  which  the  alisphenoid  wing  of  the  internal 
pterygoid  fossa  is  expanded,  and  the  alisphenoid  foramen 
enters  the  skull  through  the  medial  surface  of  the  alisphenoid. 
This  region  of  the  skull  of  Eucastor  was  not  preserved  well 
enough  to  allow  comparison. 

Dipoides  does  not  possess  the  dorsal  choana  (mesoptery- 
goid  fossa)  of  Castoroides  (see  Stirton,  1965).  This  fossa  is 
unique  to  Castoroides.  Castor  does  have  a shallow  invagi- 
nation in  the  basioccipital,  but  this  invagination  is  posterior 
to  the  mesopterygoid  fossa  of  Castoroides. 

A small  anterior  carotid  foramen  in  Dipoides  pierces  the 
lateral  margin  of  the  basisphenoid  anterior  to  the  medial 
extension  of  the  bulla.  It  is  similar  in  position  to  that  of 
Castor.  In  Dipoides  and  Eucastor,  the  posterior  carotid  fo- 
ramen is  not  visible  along  the  medial  margin  of  the  bulla 
anterior  to  the  posterior  lacerate  foramen  as  it  is  in  Castor 
and  Castoroides.  A large  posterior  lacerate  foramen  is  present 
along  the  posteromedial  border  of  the  bulla.  Directly  pos- 
terior to  the  posterior  lacerate  is  a large  hypoglossal  foramen. 
A small  stylomastoid  foramen  is  present,  wedged  between 
the  base  of  the  mastoid  process  and  the  boney  tube  of  the 
tympanic  leading  to  the  externa!  auditory  meatus.  The  par- 
occipital  process  extends  posteriorly  almost  as  far  as  do  the 
occipital  condyles  (see  Table  1).  The  very  tip  of  the  paroc- 
cipital  process  is  hooked  ventromedially. 

The  bullae  are  very  inflated.  Their  ventral  extension  does 
not  reach  as  far  ventrally  as  the  greatest  depth  of  the  skull 
at  the  tooth  row  (see  Fig.  2,  B;  Table  2).  In  Castor,  the  bulla, 
though  exhibiting  no  greater  inflation,  extends  ventrally  to  a 


Table  2.  Measurements  of  cranial  and  dental  characters  of  Dipoides 


smithi,  UCMP  113422  (in  millimeters). 

Posterior  margin  lambdoidal  crest  to  anterior 
tip  of  nasals  90.5* 

Dorsal  width  of  rostrum  anterior  to 
zygomatic  arch  I 7.0 

Least  width  between  orbital  fossae  14.7 

Greatest  width  between  external  auditory  meatus  45.7 

Greatest  width  between  mastoid  processes  23.3 

Greatest  width  of  zygomatic  arches  64.9 

Greatest  width  between  masseteric 

superficialis  processes  23.3 

Length  from  dorsal  margin  of  foramen  magnum 
to  greatest  height  of  occiput  15.2 

Depth  of  skull  between  dorsal  surface  of  skull 
and  occlusal  surface  of  M2  36.8 

Depth  of  skull  between  sagittal  crest  and  bulla  30.6 

Least  width  of  palate  between  P4’s  5.2 

Least  width  of  palate  between  M3’s  14.4 

Length  from  anterior  border  of  PJ  to  posterior 
border  of  M3  21.9 

Length  of  occlusal  surface  of  P4  5.8 

Width  of  occlusal  surface  of  P4  5.5 

Length  of  occlusal  surface  of  M1  4.9 

Width  of  occlusal  surface  of  M1  4.5 

Length  of  occlusal  surface  of  M2  5.0 

Width  of  occlusal  surface  of  M2  4.4 

Length  of  occlusal  surface  of  M3  5.1 

Width  of  occlusal  surface  of  M3  3.8 


* Approximate. 


level  in  line  with  the  tooth  row,  which  is  considerably  more 
ventral  than  in  Dipoides,  Eucastor,  or  Castoroides. 

In  ventral  view,  the  skull  of  Dipoides  closely  resembles 
those  of  Eucastor  and  Castor  in  general  outline.  The  incisors 
form  a crescentic  arc  that  terminates  above  P4.  The  dilator 
naris  fossa  is  deep  in  Dipoides,  with  a well-developed,  grooved 
median  ridge  present  on  the  maxilla.  In  Eucastor,  the  dilator 
naris  fossa  is  deep,  as  in  Dipoides ; however,  in  the  former, 
the  maxilla  slopes  steeply  toward  the  maxillary-premaxillary 
suture,  and  in  the  latter  a relatively  flat  platform  is  present. 
The  masseteric  superficialis  process  of  Dipoides  and  Casto- 
roides is  posterior  to  the  anterior  flaring  of  the  maxillary. 
The  cheek  tooth  row  diverges  posteriorly  from  the  midline 
of  the  skull.  Castoroides  has  a similar  appearance  with  closely 
placed  P4’s  and  widely  separated  M3’s;  whereas  in  Eucastor 
and  Castor,  the  cheek  tooth  rows  are  nearly  parallel.  The 
tooth  wear  in  Dipoides  forms  a flat  transverse  plane  differing 
from  that  in  Castor  wherein  the  upper  tooth  surface  is  con- 
cave with  the  labial  margin  having  greater  elevation  than  the 
lingual  edge.  There  are  two  nearly  parallel  medial  grooves 
that  run  from  the  premaxillary-maxillary  suture  posterior  to 
ML  These  grooves  resemble  those  in  Eucastor  and  Castor, 


Contributions  in  Science,  Number  346 


Wagner:  Cranial  Morphology  of  Dipoides  smithi  5 


though  in  Dipoides  they  are  deeper.  The  anterior  palatine 
foramina  are  present  on  the  suture  between  the  M2’s.  The 
posterior  palatine  foramen  is  situated  directly  posterior  to 
M3.  A ridge  is  present  posterior  to  the  posterior  palatine 
foramen  forming  the  posterolateral  margin  of  the  palate.  The 
posterior  margin  of  the  palate  forming  the  roof  of  the  mouth 
of  Dipoides  describes  a smooth  concave  margin.  Eucastor 
differs  markedly  from  this  condition  in  having  an  invagi- 
nation in  the  palatine  bone  that  extends  anteriorly  to  the 
anterior  margin  of  M3.  Castor  and  Castoroides  resemble  Di- 
poides in  this  feature. 

DISCUSSION 

A detailed  examination  of  the  cranial  morphology  of  Di- 
poides smithi  reveals  close  similarities  to  Eucastor  cf.  E. 
tortus  and  Castoroides  ohioensis.  The  similarities  in  the  po- 
sitions of  the  foramina  in  the  orbital  region  and  basicranium 
and  in  the  bone  development  of  the  skull  in  these  three 
species  indicate  a close  phyletic  relationship  supporting  Stir- 
ton’s  (1935)  conclusions  based  on  dental  morphology.  The 
similarities  of  the  skull  in  Dipoides,  Eucastor,  and  Casto- 
roides are  ( 1 ) the  relationship  of  bones  of  the  lacrimal  region; 
(2)  the  arrangement  of  foramina  in  the  basicranium  and  in- 
terorbital region;  and  (3)  the  development  of  the  alisphenoid 
wing  of  the  internal  pterygoid  fossa.  Castor  differs  markedly 
from  other  genera  in  all  three  characteristics. 

In  Dipoides  and  Castoroides,  the  dorsal  surface  of  the  lac- 
rimal region  is  composed  of  the  maxilla,  premaxilla,  and 
frontal  bones.  Eucastor  has  a lacrimal  component.  The  most 
anterior  extension  of  the  jugal  does  not  reach  the  lacrimal 
region.  The  jugal  does  contribute  to  this  region  of  the  skull 
in  Castor,  a definite  difference  from  the  other  mentioned  taxa. 

In  the  Eucastor,  Dipoides,  Castoroides  group,  Eucastor  and 
Dipoides  differ  in  the  morphology  of  the  infraorbital  foramen 
from  Eucastor  to  Dipoides.  The  structure  of  the  infraorbital 
foramen  in  Dipoides  and  Castoroides  is  derivable  from  the 
condition  found  in  Eucastor.  The  multiple  presphenoid  ca- 
nals in  Eucastor,  Dipoides,  and  some  specimens  of  Castor 
appear  to  be  a primitive  trait  evolved  prior  to  Eucastor  and 
retained  in  Eucastor,  Dipoides,  and  Castor  but  lost  in  Cas- 
troides. 

The  nonparallel  upper  cheek  tooth  rows  of  Dipoides  and 
Castoroides  are  derived  compared  with  Eucastor  and  Castor. 
Similar  nonparallel  tooth  rows  are  seen  in  some  South  Amer- 
ican histricomorph  rodents.  It  is  probable  that  this  represents 
a modification  related  to  a different  masticatory  adaptation. 
For  example,  the  South  American  rodents  that  exhibit  this 


type  of  modification  are  primarily  aquatic  forms,  with  teeth 
composed  of  lophs  for  grinding,  very  similar  to  the  castorid 
genera. 

ACKNOWLEDGMENTS 

Appreciation  is  expressed  to  the  Oregon  Museum  of  Science 
and  Industry  (OMSI)  for  providing  me  the  opportunity  to 
prospect  and  collect  fossil  specimens  from  eastern  Oregon 
during  the  summer  of  1974,  in  conjunction  with  the  Paleon- 
tological Research  Team,  during  which  time  the  specimen 
of  Dipoides  smithi  described  in  this  paper  was  collected.  All 
members  of  the  Paleontology  Research  Team,  sponsored  by 
OMSI,  deserve  credit  for  their  participation,  sharp  eyes,  and 
enthusiasm  during  the  field  season.  It  was  a pleasure  having 
Bruce  Batten,  Steve  Damiani,  Kathy  Harvey,  Roy  West,  and 
Kathy  Wolfram  as  students  and  companions  during  the  six- 
week  program.  The  near  perfectly  preserved  skull  of  Dio- 
poides  smithi  recovered  from  McKay  Reservoir  was  found 
by  Bruce  Batten.  The  illustrations  of  D.  smithi  were  drawn 
by  Jaime  P.  Lufkin,  an  artist  from  the  Museum  of  Paleon- 
tology, University  of  California,  Berkeley. 

I thank  the  curators  of  the  Museum  of  Paleontology,  Uni- 
versity of  California,  Berkeley,  The  University  of  California 
Museum  of  Vertebrate  Zoology,  and  the  Natural  History 
Museum  of  Los  Angeles  County  for  the  loan  and  use  of 
comparative  collections.  I also  thank  Lawrence  G.  Barnes 
for  the  helpful  and  constructive  comments  offered  in  his 
review  of  this  manuscript. 

LITERATURE  CITED 

Jager,  G.F.  1835.  Ueber  die  fossilen  Saugethiere,  welche 
in  Wtirttemberg  gefunded  worden  sind.  Le  Abth;  17-18. 
Olson,  E.C.  1940.  Cranial  foramina  of  North  American 
beavers.  Jour.  Paleontology  14:495-501. 

Shotwell,  J.A.  1955.  Review  of  the  Pliocene  beaver  Di- 
poides. Jour.  Paleontology  29:129-144. 

Stirton,  R.A.  1935.  A review  of  the  Tertiary  beavers.  Univ. 
Ca.  Pubs.,  Geol.  Sci.,  Bull.  23:391-458. 

. 1965.  Cranial  morphology  of  Castoroides.  Dr.  D.N. 

Wadia  Comm.  Vol.  Nim.  & Metal.  Inst,  of  India,  pp. 
273-285. 

Wagner,  H.M.  1981.  Geochronology  of  the  Mehrten  For- 
mation in  Stanislaus  County,  California.  Ph.D.  disser- 
tation, Dept.  Earth  Sci.,  Univ.  of  California,  Riverside, 
347  pp. 

Submitted  22  June  1982;  accepted  3 May  1983. 


6 Contributions  in  Science,  Number  346 


Wagner:  Cranial  Morphology  of  Dipoides  smithi 


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SCIENTIFIC  PUBLIC.  ATHENS  COMMITTEE 

'Craig  C.  Black  Museum  Director 
Donald  Output 

Daniel  M.  Cohen,  Committee  Chairman 
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Robin  A.  Simpson,  Managing  Editor 
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IAR  2 8 1984 
Lib  rabies 


STUDIES  ON  THE  TAXONOMY  AND  DISTRIBUTION  OF 
AMERICAN  CENTRIDINE  BEES 
(HYMENOPTERA:  ANTHOPHORIDAE) 

Roy  R.  Snelling1 


ABSTRACT.  This  study  focuses  primarily  on  the  Centridini  of 
North  America  south  of  the  United  States  and  includes  a key  to  the 
three  genera  of  Centridini:  Centris,  Ptilotopus,  and  Epicharis.  Within 
the  genus  Centris  keys  are  provided  for  species  in  the  subgenera 
Paracentris,  Centris,  Xanthemisia,  Acritocentris,  Melanocentris, 
Trachina,  Hemisiella,  and  Heterocentris ; one  new  monotypic  sub- 
genus is  described:  Ptilocentris (type-species:  Centris festiva  F.  Smith, 
1854). 

The  following  new  species  are  described:  C.  ( Xerocentris ) griseola 
(Mexico);  C.  (C.)  aethiocesta  (El  Salvador  to  Panama);  C.  (Acrito- 
centris) satana  (Mexico,  United  States);  C.  (Melanocentris)  agi/oides 
(Mexico  to  Costa  Rica);  C.  (M.)  gelida  (Mexico,  Guatemala);  C. 
(Trachina)  eurypatana  (Mexico);  C.  (T.)  xochipillii (Mexico).  Centris 
(C.)  meaculpa  is  proposed  as  a new  name  for  C.  (C.)  erubescens 
Snelling,  1974,  not  C.  costaricensis  var.  erubescens  Friese,  1925. 
Centris  atripes  var.  ferrisi  Cockerell  is  a valid  species  in  Paracentris ; 
the  types  are  males,  not  females  as  stated  by  Cockerell. 

The  following  new  synonymy  is  proposed  in  Centris:  C.  cockerelli 
resoluta  Cockerell  = C.  (Paracentris)  cockerelli  W.  Fox;  C.  clypeata 
Friese,  C.  anthracina  Snelling  = C.  (P.)  nigrocaerulea  F.  Smith;  C. 
limbata  Friese  = C.  ( P .)  atripes  Mocsary;  C.  strawi  Snelling  = C. 
(Acritocentris)  albiceps  Friese;  C.  robusta  Cockerell,  C.  segregata 
Crawford  = C.  (C.)  inermis  Friese;  C.  citrotaeniata  Gribodo,  C.Jla- 
vifrons  var.  rufescens  Friese,  C.  flavifrons  var.  nigritula  Friese  = C. 
(C.)  flavifrons  (Fabricius);  C.  chlorura  Cockerell  = C.  (Ptilocentris) 
festiva  F.  Smith;  C.  ignita  F.  Smith,  C.  bakerella  Friese,  Epicharis 
cisnerosi  Cockerell  = C.  (Melanocentris)  agilis  F.  Smith;  C.  fusci- 
ventris  var.  scutellata  Mocsary  = C.  (M.)  fusciventris  Mocsary;  C. 
melanochlaena  F.  Smith,  Epicharis  zamoranensis  Cockerell  = C.  (A/.) 
obsoleta  Lepeletier;  C.  schwarzi  Cockerell  = C.  (Trachina)  labiata 
Friese;  C.  confinis  Perez  = C.  (Hemisiella)  nitida  F.  Smith;  C.  ruae 
Cockerell  = C.  (H.)  transversa  Perez;  C.  dentipes  F.  Smith,  C.  ru- 
fomaculata  Cockerell,  C.  lanipes  subtarsata  Cockerell  = C.  (H.)  tri- 
gonoides  Lepeletier;  C.  costaricensis  var.  erubescens  Friese  = C.  (H.) 
vittata  Lepeletier;  C.  triangulifera  Cockerell  = C.  (Heterocentris) 
labrosa  Friese. 

Ptilotopus  is  elevated  to  generic  rank  from  its  previous  status  as 
a subgenus  of  Centris ; P.  zonalis  Mocsary  of  Panama  is  the  only 
species  in  North  America. 

A key  is  given  for  all  the  subgenera  of  Epicharis  and  keys  are  given 
for  the  species  of  each  subgenus  known  to  occur  in  North  America. 


Two  new  species  are  described  in  the  subgenus  Epicharana:  E.  an- 
gulosa  (Costa  Rica)  and  E.  bova  (Costa  Rica,  Panama).  The  following 
are  new  synonyms:  E.  salazari  Cockerell  = E.  (Epicharana)  elegans 
F.  Smith;  E.  rustica  var . /lava  Friese  = E.  (Epicharana)  rustica  (Oliv- 
ier); E.  phenacura  Cockerell,  E.  conura  Cockerell  = E.  (Parepicharis) 
metatarsalis  Friese.  North  American  distribution  data  are  cited  for 
all  species. 

Included  is  a synonymic  list  of  the  recognized  North  and  Central 
American  Centridini. 

RESUMEN.  Este  estudio  se  refiere  principalmente  a los  Centridini 
de  Norte  America  al  sur  de  los  Estados  Unidos  e incluye  una  clave 
para  los  tres  generos  de  Centridini:  Centris,  Ptilotopus  y Epicharis. 
Dentro  del  genero  Centris  se  incluyen  claves  para  ias  especies  de  los 
subgeneros  Paracentris,  Centris,  Xanthemisia,  Acritocentris.  Mela- 
nocentris, Trachina,  Hemisiella  y Heterocentris ; un  subgenero  mo- 
notipico  nuevo  se  describe,  Ptilocentris  (especie-tipo:  Centris  festiva 
F.  Smith,  1854). 

Ptilotopus  se  eleva  al  rango  generico  desde  su  estatus  previo  de 
subgenero  de  Centris ; P zonalis  Mocsary  de  Panama  es  la  unica 
especie  en  Norte  America. 

Se  da  una  clave  para  todos  los  subgeneros  de  Epicharis  y para  las 
especies  de  cada  subgeneros  conocidos  en  Norte  America. 

Se  describen  ocho  nuevas  especies  para  el  genero  Centris  y dos 
para  el  genero  Epicharis.  Estas  al  igual  que  los  nuevos  sinommos  se 
incluyen  en  el  “Abstract.” 

Se  citan  datos  de  distribucion  de  Norte  America  para  todas  las 
especies.  Se  incluye  una  lista  de  los  Centridini  reconocidos  de  Norte 
America  y de  las  sinonimias  conocidas. 

INTRODUCTION 

The  present  paper  is  a continuation  of  my  earlier  work  on 
the  Centris  of  North  and  Central  America  (Snelling,  1956, 
1 966,  1974).  New  data  on  the  distribution  of  previously  treat- 
ed species,  as  well  as  investigations  into  the  systematics  of 
previously  unstudied  groups  are  presented  below.  Although 
my  previous  investigations  involved  only  the  genus  Centris, 


1.  Section  of  Entomology,  Natural  History  Museum  of  Los  An- 
geles County,  900  Exposition  Blvd.,  Los  Angeles,  California  90007. 


Contributions  in  Science,  Number  347,  pp.  1-69 
Natural  History  Museum  of  Los  Angeles  County,  1984 


ISSN  0459-8113 


the  study  has  now  expanded  to  include  the  genera  Epicharis 
and  Ptilotopus,  the  latter  newly  elevated  to  generic  status 
from  its  previous  position  as  a subgenus  of  Centris. 

SPECIMENS  EXAMINED 

Material  utilized  in  this  study  is  from  the  following  institu- 
tional and  private  collections:  American  Museum  of  Natural 
' History  (AMNH),  Robert  W.  Brooks,  personal  collection 
’ (RWB),  California  Academy  of  Sciences  (CAS),  Cornell  Uni- 
versity (CORN),  Florida  State  Department  of  Plant  Industry 
(DPIF),  Museum  of  Comparative  Zoology  (MCZ),  Museum 
National  d’Histoire  Naturelle,  Paris  (MNHN),  Natural  His- 
tory Museum  of  Los  Angeles  County  (LACM),  John  L.  Neff, 
personal  collection  (NEFF),  Oregon  State  University  (ORSU), 
D.  Roubik,  personal  collection  (ROUB),  United  States  Na- 
tional Museum  of  Natural  History  (USNM),  University  of 
California,  Berkeley  (UCB),  University  of  California,  Davis 
(UCD),  University  of  Kansas  (UKAN),  and  Thomas  J.  Za- 
vortink,  personal  collection  (TJZ). 

SPECIMEN  DATA 

New  data  on  distribution,  capture  dates,  and  plant  associa- 
tion are  fully  cited  for  many  species  since  there  are  few  pub- 
lished records.  Localities  are  organized  by  country  and  by 
state,  province  or  department  within  that  country.  The  des- 
ignation “state,”  “department,”  or  “province”  is  omitted  to 
conserve  space.  Thus,  a locality  cited  under  “COSTA  RICA, 
SAN  JOSE”  is  from  San  Jose  Province  in  Costa  Rica. 

Specimen  data  are  cited  for  most  species,  but  some  species 
are  common,  widely  distributed  and  abundant.  For  such 
species,  only  general  range  is  given. 

TERMINOLOGY 

In  general,  the  morphological  terminology  employed  below 
is  that  which  is  traditional  is  apoid  systematics,  following 
Michener  (1944,  1954,  1 965).  A few  terms,  however,  should 
be  explained,  in  order  that  they  may  be  correctly  employed 
(in  the  sense  of  this  paper)  by  others. 

Antennal  socket  diameter.  Since  the  antennal  socket  is  not 
circular  it  follows  that  two  different  measurements  are  pos- 
sible. In  this  study,  the  antennal  socket  diameter  is  measured 
perpendicular  to  the  long  axis  of  the  head  (i.e.,  the  transverse 
diameter);  measurement  is  made  from  the  summit  of  the  rim 
on  the  opposite  side.  Associated  measurements  include  the 
interantenna/  distance  and  the  antennocular  distance. 

Interantenna/  distance  is  measured  as  the  least  distance 
between  the  summits  of  the  two  antennal  socket  rims. 

Antennocular  distance  is  the  least  distance  from  the  inner 
eye  margin  to  the  summit  of  the  rim  of  the  adjacent  antennal 
socket. 

Several  features  of  the  clypeus  should  be  defined  for  con- 
sistent usage.  The  clypeal  disc  is  the  central  portion  of  the 
clypeus,  from  base  to  apical  margin  and  roughly  bounded  on 
either  side  by  an  imaginary  line  extended  distad  from  the 
junction  of  the  subantennal  sutures  with  the  base  of  the  clyp- 


eus. Clypeal  length,  when  compared  to  clypeal  width,  is  mea- 
sured along  the  midline,  from  the  base  to  the  apical  margin. 
Clypeal  width  is  measured  as  the  greatest  distance  between 
the  apicolateral  extremities  of  the  clypeus  (i.e.,  that  portion 
nearest  the  inner  eye  margin).  The  distance  between  the  clyp- 
eus and  the  eye  is  measured  as  the  least  distance  between  the 
apicolateral  extremity  and  the  nearest  point  on  the  inner  eye 
margin. 

Elaiospathe  is  a new  term:  It  is  a combination  of  the  Greek 
words  elaion  (oil)  and  spathe  (a  paddle  for  stirring  or  mixing; 
a broad  blade).  This  term  refers  to  the  modified  oil-gathering 
structures  present  in  female  centridine  and  exomalopsine 
bees.  The  elaiospathe  most  commonly  consists  of  a row  of 
long,  erect,  apically  spatulate  and  curved  setae,  usually  on 
the  pro-  and  mesobasitarsi;  in  Centridini  they  are  situated 
anteriorly  on  the  “inner”  surface  of  the  segment,  in  some 
Exomalopsini  they  are  external  and  may  be  present  in  males 
as  well  as  females,  and  are  limited  to  the  probasitarsi.  How- 
ever, oil-collecting  structures  may  also  be  present  on  the 
abdominal  venter,  at  least  in  some  species  of  Tapinotaspis 
( Tapinotaspoides).  For  an  excellent  survey  of  the  morphology 
and  function  of  the  elaiospathe,  see  Neff  and  Simpson  (198 1). 

The  lower  frontal  width  is  measured  as  the  distance  be- 
tween the  inner  eye  margins  at  the  level  of  the  apicolateral 
angle  of  the  clypeus.  It  is  compared  with  the  upper  frontal 
width  to  determine  the  degree  of  convergence  of  the  inner 
eye  margins.  The  upper  frontal  width  is  the  minimum  dis- 
tance between  the  inner  eye  margins  at  about  the  level  of  the 
ocelli. 

The  diameter  of  the  anterior  ocellus  is  the  greatest  trans- 
verse (perpendicular  to  long  axis  of  head)  diameter  of  the 
anterior  ocellus.  Interocellar  distance  is  the  minimum  dis- 
tance between  the  posterior,  or  lateral,  ocelli.  Ocellocular 
distance  is  the  minimum  distance  between  one  of  the  pos- 
terior ocelli  and  the  nearest  point  on  the  eye  margin.  Ocel- 
loccipital  distance  is  measured  in  dorsal  view  and  is  the  min- 
imum distance  from  the  posterior  margin  of  one  of  the 
posterior  ocelli  to  the  occipital  declivity.  Transocellar  dis- 
tance is  measured  as  the  maximum  distance  between  the 
outer  margins  to  the  two  posterior  ocelli.  In  all  cases,  the 
ocellus  is,  for  purposes  of  measurement,  considered  to  be 
only  the  transparent  amber-colored  lens. 

Pilosity.  In  general,  centridines,  like  most  other  anthopho- 
rids,  are  densely  hairy  bees,  with  much  of  the  head,  thorax, 
and  first  abdominal  tergum  clothed  with  long,  plumose,  de- 
cumbent to  fully  erect  hairs.  Similarly,  the  legs  are  hairy,  but 
hairs  tend  to  be  decumbent  to  subappressed  on  the  outer 
faces  of  the  tibiae  and  basitarsi  and  are  often  stout  and  simple. 
Long,  plumose  hairs  are  usually  present,  but  sparse,  on  the 
femora.  On  the  abdomen,  the  hairs  on  the  discs  of  the  seg- 
ments generally  become  longer  and  more  erect  on  succeeding 
segments,  plumose  hairs  as  a rule  being  absent  from  the  discs 
of  the  second  and  third  terga. 

The  abdominal  sterna  normally  have  long,  plumose  hairs 
which  tend  to  be  longest  and  most  abundant  along  the  mid- 
line. As  a rule,  males  are  more  densely  pubescent  on  the 
sterna  than  are  the  females. 

Punctation.  The  description  of  characteristics  related  to 


2 Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


the  punctation  of  the  integument  of  bees  and  other  Hyme- 
noptera  have  long  been  the  bane  of  taxonomists  since  the 
terminology  has  always  been  undefined  and  subject  to  con- 
siderable variation  in  interpretation.  Punctures  which  one 
author  considers  to  be  fine,  another  might  describe  as  mod- 
erate in  size.  The  distances  between  punctures,  whether  dense, 
close,  or  sparse,  have  likewise  been  troublesome.  A few  years 
ago  I (Snelling,  1980)  introduced  a system  of  terminology  to 
solve  this  difficulty  in  the  genus  Hylaeus.  After  considering 
the  objections  by  Harris  (1979),  I can  find  no  valid  reason 
not  to  apply  the  same  terminology  of  absolute  measurements 
to  the  Centridini.  I do  not  agree  that  punctation  need  be 
described  relative  to  the  size  of  the  entity.  Admittedly,  I was 
initially  uncomfortable  at  describing  as  “coarse”  those  punc- 
tures on  a centridine  bee  that  I was  accustomed  to  considering 
to  be  moderate-sized.  This  is,  however,  strictly  a perceptual 
problem,  one  which  has  ceased  to  disturb  my  sense  of  “right- 
ness.” 

Therefore,  I propose  to  expand  the  usage  of  my  system  to 
include  the  Centridini.  Such  a system  must  be  considered, 
at  this  point,  to  be  experimental;  some  adjustment  or  mod- 
ification may  prove  to  be  necessary  as  this  method  is  tried 
on  other  groups.  A micrometer  disk  in  one  ocular  of  a mi- 
croscope is,  obviously,  necessary  and  I use  a minimum  mag- 
nification of  64  x;  several  punctures  of  about  the  same  ap- 
parent size  are  measured  and  averaged.  The  appropriate 
terminology  is  as  follows: 

minute— puncture  diameter  0.010-0.019  mm 
fine— puncture  diameter  0.020-0.035  mm 
moderate— puncture  diameter  0.036-0.055  mm 
coarse— puncture  diameter  0.056-0.070  mm 
very  coarse  — puncture  diameter  over  0.070  mm 

Since  punctures  are  often  not  uniform  in  size  on  a given 
segment  or  area,  it  may  be  necessary  to  combine  terms  into 
phrases  such  as  “fine  to  moderate”  (puncture  diameter  vary- 
ing between  0.020  and  0.055  mm),  though  usually  a more 
limited  size  range,  such  as  moderate,  may  prevail. 

The  relative  density  of  punctures  may  also  be  standardized 
and  I use  the  following  terminology  below: 

Contiguous— punctures  so  close  that  they  are  often  de- 
formed; interspaces  are  compressed  and  sharp-edged. 

Subcontiguous— punctures  separated  by  more  or  less  flat 
interspaces  up  to  about  0.30  times  a puncture  diameter;  some 
punctures  may  be  deformed. 

Dense  — punctures  separated  by  more  or  less  flat  inter- 
spaces between  0.30  and  0.70  times  a puncture  diameter; 
punctures  usually  round  but  may  be  elongate. 

Close  — punctures  separated  by  more  or  less  flat  interspaces 
0.70  to  1.50  times  a puncture  diameter. 

Sparse  — punctures  separated  by  more  or  less  flat  inter- 
spaces 1.50  to  3.00  times  a puncture  diameter. 

Scattered — puncture  interspaces  are  very  irregular  and  range 
from  about  3.00  to  6.00  or  more  times  a puncture  diameter. 

Variations  in  puncture  density  may  be  expressed  by  com- 
bining terms:  “sparse  to  scattered.” 

Clypeal  punctation  is  usually  described  from  the  middle 
one-third  of  the  disc. 


Mesoscutal  punctation  is  described  from  the  area  between 
the  parapsidal  line  and  the  median  line  at  the  level  of  the 
tegula;  punctures  are  commonly  less  close  posteromesally  on 
the  segment  and  are  closer  laterad  of  the  parapsidal  line. 
Scutellar  and  metanotal  sculpture  is  described  from  the  mesal 
one-third  of  the  segment.  The  middle  of  the  mesopleural  disc 
is  the  standard  for  that  segment. 

The  first  abdominal  segment  has  a nearly  vertical  anterior 
and  a horizontal  posterior  face;  the  latter  is  referred  to  as  the 
disc  of  the  segment.  Tergal  punctation  is  described  from  the 
mesal  one-third  of  the  postgradular  area  of  the  segments 
beyond  the  first,  and  anterior  to  the  apical  zone. 

The  distal  portion  of  the  abdominal  terga,  especially  of  the 
second  and  following  segments,  except  the  last,  is  commonly 
depressed  in  bees;  this  area  is  usually  called  the  apical  depres- 
sion and  it  is  often  sculptured  differently  from  the  disc.  In 
centridines  this  area  is  usually  not  obviously  depressed,  es- 
pecially across  the  middle  two-thirds  of  the  segment,  but  is 
usually  more  sparsely  and  less  coarsely  sculptured  than  the 
disc;  herein  I refer  to  this  as  the  apical  zone. 

In  the  description  of  new  taxa,  measurements  and  pro- 
portions are  based  on  the  primary  type  specimens.  The  cor- 
responding range  of  measurements  and  proportions  of  para- 
types  only  (if  any)  are  indicated  parenthetically  after  those 
of  the  primary  type. 

NESTING  BIOLOGY 

The  little  that  is  known  of  centridine  nesting  biology  was 
reviewed  by  Coville,  Frankie,  and  Vinson  (1983).  They  con- 
cluded that  the  choice  of  nesting  substrate  by  the  female  bees 
is  roughly  correlated  with  taxonomic  groupings.  Thus,  those 
species  in  the  subgenera  Wagenknechtia,  Paracentris,  Xero- 
centris,  Centris  s.  str.,  Exallocentris,  and  Melanocentris  nor- 
mally excavate  original  tunnels  in  soil.  About  one-half  of  the 
species  cited  utilize  flat  soil  and  about  one-half  excavate  in 
more  or  less  vertical  banks  or  mud  walls;  two  species  evi- 
dently are  capable  of  choosing  either  flat  or  vertical  surfaces. 

Species  belonging  to  the  subgenera  Hemisiella  and  Het- 
erocentris  generally  utilize  preexisting  cavities  in  a variety  of 
substrates.  Some  species,  at  least,  seem  to  be  highly  oppor- 
tunistic in  choice  of  substrate  and  may  choose  old  bee  cells 
and  burrows  in  soil,  abandoned  Sceliphron  nests,  or  holes  in 
wood.  Of  two  species  of  Trachina  cited,  one  nested  in  the 
walls  of  a living  arboreal  termitarium  and  the  other  chose 
flat  soil. 

Species  of  Ptilotopus,  herein  treated  as  a separate  genus, 
were  uniform  in  excavating  their  nests  in  the  walls  of  living 
arboreal  termitaria.  There  is  some  suggestion  that  the  ar- 
rangement of  cells  within  the  nest  may  similarly  substantiate 
the  present  groupings  of  subgenera,  but  the  data  are  very 
fragmentary. 

SYSTEM  A I ICS 

The  general  interpretation  of  the  Centridini  has  been  that 
there  are  two  genera,  Centris  and  Epicharis  (e.g.,  Michener, 
1954).  One  result  of  my  recent  work  in  this  tribe  is  that  I 
now  believe  that  Ptilotopus , formerly  treated  as  a subgenus 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  3 


of  Centris,  must  be  elevated  to  generic  status;  justification 
for  this  decision  will  be  provided  below.  The  three  genera  of 
Centridini  which  1 recognize  may  be  separated  as  follows. 

KEY  TO  GENERA  OF  CENTRIDINI 

la.  Marginal  cell  of  forewing  shorter  than  distance  from  its 
apex  to  tip  of  wing;  occipital  llagelliform  setae  usually 
absent,  but  if  present,  not  reaching  as  far  back  as  anterior 

margin  of  tegula 2 

b.  Marginal  cell  of  forewing  longer  than  distance  from  its 
apex  to  tip  of  wing;  occipital  llagelliform  setae  present 
and  usually  extending  to,  or  beyond,  level  of  anterior 

margin  of  tegula Epicharis 

2a.  Occipital  llagelliform  setae  absent;  mesosternum  of  fe- 
male without  tubercles;  hypoepimeron  flat  or  slightly 
convex;  pronotal  lobe  low  and  moderately  convex,  never 

conspicuously  tuberculate  Centris 

b.  Occipital  llagelliform  setae  present  and  extending  to  an- 
terior margin  of  mesoscutum,  or  slightly  beyond;  meso- 
sternum of  female  with  a pair  of  stout  tubercles  hidden 
in  dense  cluster  of  hairs;  hypoepimeron  usually  promi- 
nently coniform,  but  if  not,  its  lower  margin  is  sharply 
elevated  above  adjacent  portions  of  mesepisternum;  pro- 
notal lobe  often  tuberculate Ptilotopus 

Genus  Centris  Fabricius 

Centris  is  a primarily  tropical  genus,  with  1 1 subgenera;  these 
subgenera  are  separated  in  keys  by  Michener  (1951)  and 
Snelling  (1974).  The  separation  of  Ptilotopus  from  Centris 
(see  below)  removes  the  only  group  with  flagelliform  occipital 
setae.  The  North  American  species  of  the  subgenera  Xero- 
centris  and  Paracentris  were  treated  by  Snelling  (1974).  The 
North  American  species  of  the  remaining  subgenera  are  treat- 
ed below. 

Subgenus  Xerocentris  Snelling 

Centris  subg.  Xerocentris  Snelling,  1974:3-4.  Type-species; 

Centris  californica  Timberlake,  1 940;  original  designation. 

This  subgenus  was  proposed  for  a number  of  species  found 
in  arid  regions  of  North  and  South  America.  Although  Xero- 
centris is  related  to  Paracentris,  females  lack  an  elaiospathe 
on  the  pro-  and  mesobasitarsi,  present  in  Paracentris  and 
Centris  s.s.,  and  the  pygidial  plate  is  convex  along  the  median 
line.  Males  are  much  like  those  of  Paracentris.  but  the  ocel- 
locular  distance  is  less  than  the  diameter  of  a lateral  ocellus, 
and  the  abdominal  terga  are  covered  with  abundant  fully 
erect,  often  pale,  hairs. 

One  new  species  of  Xerocentris  is  described  below. 

Centris  ( Xerocentris ) griseoia,  new  species 

DIAGNOSIS 

Female  runs  to  C.  vanduzeei  Cockerell  in  my  key  (1974),  but 
mandible  tndentate  (quadridentate  in  C.  vanduzeei).  meso- 
scutum and  scutellum  with  mixed  black  and  pale  erect  hairs 
(wholly  pale  in  C.  vanduzeei ),  prepygidial  brush  blackish 


brown  (bright  fulvous  in  C.  vanduzeei)  and  distal  portion  of 
median  ridge  of  pygidium  low  and  rounded  (cariniform  in 
C.  vanduzeei).  Male  unknown. 

DESCRIPTION 

HOEOTYPE  FEMALE.  Measurements  (mm).  Head  width 
5.18  (5.03-5.28);  head  length  3.79  (3.23-3.33)  wing  length 
10.1  (10.2-10.3);  total  length  13.0  (13.0-15.5). 

Head.  1.58  (1 .5 1-1 .62)  times  broader  than  long;  occipital 
margin  nearly  Hat  in  frontal  view;  inner  orbits  strongly  con- 
vergent above,  upper  frontal  width  0.84  (0.82-0.84)  times 
lower  frontal  width.  Maxillary  palp  five-segmented.  Man- 
dible slender,  tndentate,  second  preapical  tooth  absent;  inner 
tooth  prominent,  triangular,  and  nearly  perpendicular  to  long 
axis  of  mandible.  Labrum  unmodified;  shiny  between  sub- 
contiguous,  moderate  punctures.  Clypeus  flattened,  weakly 
protuberant;  disc  slightly  shiny,  surface  roughened,  irregu- 
larly, obliquely  to  transversely  rugose.  Frons  and  vertex  mod- 
erately shiny  to  shiny  between  fine  subcontiguous  to  dense 
punctures,  but  with  extensive  smooth  areas  in  front  of,  and 
laterad  to,  ocelli;  gena  less  than  half  as  wide  as  eye,  mod- 
erately shiny  between  fine,  dense  punctures.  Interantennal 
distance  2.92  (2.32-3.67)  times  antennal  socket  diameter; 
antennocular  distance  1.62(1 .65-2.00)  times  antennal  socket 
diameter;  scape  short,  robust,  scape  length  0.73  (0.67-0.70) 
times  length  of  first  flagellar  segment;  first  flagellar  segment 
4.83  (5.06-5.50)  times  length  of  second.  Ocelli,  in  frontal 
view,  well  below  occipital  margin;  interocellar  distance  2.23 
(2.20-2.24)  times  ocellar  diameter;  ocellocular  distance  1 .50 
(1.52-1.58)  times  ocellar  diameter;  ocelloccipital  distance 
2.50  (2.52-2.65)  times  ocellar  diameter. 

Thorax.  Mesoscutum  shiny  between  subcontiguous  to  dense 
moderate  punctures;  scutellum  narrowly  shiny  and  impunc- 
tate  across  anterior  margin  but  mostly  weakly  shiny,  inter- 
spaces conspicuously  roughened  between  dense,  moderate 
punctures;  metanotum  vertical,  shiny  between  sparse,  fine 
punctures;  mesopleuron  moderately  shiny,  moderately  tes- 
sellate  between  subcontiguous  to  dense,  fine  punctures;  meta- 
pleuron  similar  but  a little  shinier.  Propodeal  disc  smooth 
and  shiny  between  scattered  minute  to  fine  punctures;  side 
of  propodeum  less  shiny,  punctures  dense  to  sparse,  minute. 
Metabasitibial  plate  about  twice  longer  than  broad,  strongly 
narrowed  apicad,  secondary  plate  poorly  defined  and  merging 
into  primary  plate,  anterior  depression  small  (all  specimens 
worn,  details  uncertain);  scopal  hairs  largely  plumose  along 
margins,  mostly  simple  elsewhere. 

Abdomen.  Discs  of  all  terga  moderately  shiny  and  weakly 
tessellate  between  dense  to  sparse,  fine  punctures;  pygidium 
elongate-triangular,  apex  narrowly  rounded;  median  eleva- 
tion rounded  and  not  attaining  apex. 

Color.  Generally  blackish  brown,  abdominal  terga  with 
weak  blue-green  reflections;  mandibles  dark  ferruginous;  fla- 
gellum and  legs  brown;  tergal  margins  yellowish;  tegula  pi- 
ceous;  wings  slightly  brownish,  veins  and  stigma  brown. 

Pilosity.  Of  head,  thorax  and  abdomen  fully  erect,  whitish, 
with  that  of  vertex  largely  brownish;  pale  and  blackish  hairs 
about  equally  abundant  on  mesoscutum,  scutellum  and  pro- 


4 Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


notal  lobes;  tergal  hairs  shorter  and  sparser,  plumose;  fourth 
tergite  with  a few,  and  fifth  with  many,  long,  curled  blackish 
bristles;  prepygidial  fimbria  dark;  sixth  tergite  with  dark  bris- 
tles and  hairs;  sternites  with  dark  discal  hairs  and  pale  distal 
hair  bands,  those  of  fourth  and  fifth  segments  broadly  inter- 
rupted; sixth  segment  dark  pubescent;  scopal  hairs  white; 
some  brownish  bristles  on  protibia;  bristles  of  protarsus  and 
thoracic  venter  brown;  bristles  on  inner  side  of  meso-  and 
metabasitarsi  brownish  ferruginous. 

TYPE  MATERIAL 

Holotype  female  and  three  paratypes.  MEXICO,  GUER- 
RERO: Iguala,  2000  ft.  elev.,  4 Feb.  1954  (R.R.  Dreisbach). 
Holotype  and  two  paratypes  in  UKAN;  one  paratype  in 
LACM. 

ETYMOLOGY 

Modern  Latin  griseolus  (gray)  because  of  the  grayish  ap- 
pearance of  the  thoracic  dorsum. 

DISCUSSION 

Although  C.  griseola  superficially  resembles  C.  vanduzeei 
Cockerell  of  Lower  California,  it  is  easily  separated  from  that 
species  by  the  characters  given  above  in  the  Diagnosis.  Es- 
pecially distinctive  is  the  tridentate  mandible.  The  only  other 
species  of  Xerocentris  with  tridentate  mandible  is  C.  pallida 
W.  Fox,  of  the  arid  western  deserts.  In  that  species,  the  max- 
illary palp  is  four-segmented,  the  pygidial  apex  is  truncate 
and  with  a broad  median  ridge  which  extends  to  the  apex, 
the  thoracic  integument  is  dull,  and  the  scopal  hairs  are  all 
plumose. 

The  unknown  male  of  C.  griseola  probably  resembles  the 
female  in  that  the  clypeus  will  be  black  and  the  thoracic  hairs 
will  be  a mixture  of  dark  and  light  hairs. 

Pollen  samples  from  two  specimens  were  examined  by  J.L. 
Neff.  His  conclusion  (personal  communication)  was  that  in 
both  cases,  the  samples  were  “.  . . a mixture  of  Cercidium 
and  Hoffmanseggia  (or  possibly  Caesalpinia)  with  the  former 
predominating.” 

Subgenus  Paracentris  Cameron 

Paracentris  Cameron,  1903:235-236.  Type-species:  Para- 
centris fulvohirta  Cameron,  1903;  original  designation. 
Penthemisia  Moure,  1950:390.  Type-species:  Centris  chilen- 
sis  Spinola,  1851;  original  designation. 

Hemisia,  subg.  Penthemisia:  Michener,  1951:3-4. 

Centris,  subg.  Trichocentris  Snelling,  1956:3.  Type-species: 
Centris  rhodoleuca  Cockerell,  1923;  original  designation. 
Centris,  subg.  Paracentris:  Snelling,  1974:5-7. 

The  North  American  species  of  Paracentris  were  treated  in 
1974;  14  species  were  recognized,  four  of  which  were  known 
from  only  one  sex.  Since  then,  I have  seen  opposite  sexes  for 
three  of  these  species.  One  previously  described  form,  C. 
atripes  ferrisi  Cockerell,  was  not  considered  in  1974  because 
it  was  known  only  from  the  deficient  original  description.  I 


have  now  seen  the  type  specimens.  It  seems  appropriate, 
therefore,  to  present  a new  key  to  the  North  American  species 
of  Paracentris  to  accommodate  these  new  data. 

KEY  TO  NORTH  AMERICAN  PARACENTRIS 

la.  Female,  antenna  12-segmented  and  basitibial  plate 

present  on  metatibia  2 

b.  Male,  antenna  1 3-segmented  and  basitibial  plate  absent 

from  metatibia  15 

2a.  Pubescence  of  head,  thorax,  and  legs  entirely  or  pre- 
dominantly black,  that  of  mesepistemum  wholly  black 

3 

b.  Pubescence  of  head,  thorax,  and  legs  entirely  or  pre- 
dominantly pale,  that  of  mesepistemum  pale,  at  least 


in  part  7 

3a.  Pubescence  of  thoracic  dorsum  entirely  black 4 

b.  Pubescence  of  thoracic  dorsum  largely  pale 6 


4a.  Integument  of  abdominal  tergites  black;  punctures  of 
second  tergite  uniformly  distributed  across  disc,  not 
notably  sparser  along  midline,  their  hairs  simple  or 

plumose,  interspaces  polished  or  roughened  5 

b.  Integument  of  abdominal  tergites  dark  blue;  punctures 
of  second  tergite  dense  at  sides,  distinctly  more  sepa- 
rated (sometimes  sparse)  in  middle,  their  hairs  decum- 
bent and  simple,  interspaces  polished 

laevibullata  Snelling 

5a.  Punctures  of  disc  of  second  tergite  fine,  little  greater  in 
diameter  than  hairs  arising  from  them,  hairs  fine,  plu- 
mose, suberect;  interspaces  of  second  tergite  roughened 

and  dull  nigrocaeru/ea  F.  Smith 

b.  Punctures  of  second  tergite  conspicuously  greater  in 
diameter  than  coarse,  simple,  decumbent  hairs  arising 
from  them;  interspaces  of  second  tergite  polished  and 

shiny  aterrima  F.  Smith 

6a.  Large  species,  14.5-18.5  mm  long;  pubescence  of  ver- 
tex and  pronotal  lobes  black;  clypeal  punctures  sparse, 
obscure  apicad,  median  impunctate  line  evanescent 

mexicana  F.  Smith 

b.  Smaller  species,  12.5-14.5  mm  long;  pubescence  of  ver- 
tex and  pronotal  lobe  whitish;  clypeus  with  sharply 
defined  median  impunctate  line,  punctures  sharply  de- 
fined   zacateca  Snelling 

7a.  Clypeal  integument  entirely  blackish  8 

b.  Clypeal  integument  at  least  partially  yellowish,  orange 

or  red  1 1 

8a.  Discs  of  second  and  third  terga  with  evenly  spaced, 

distinct  punctures  throughout 9 

b.  Discs  of  second  and  third  terga  polished  and  shiny, 
with  scattered  fine  punctures  which  become  coarse  and 
distinct  laterad,  most  of  disc  virtually  impunctate  . . . 

fisheri  Snelling 

9a.  First  flagellar  segment  a little  longer  than  following  three 
segments  combined;  abdominal  terga  black;  punctures 
of  scutellum  separated  by  much  less  than  a puncture 
diameter  10 

b.  First  flagellar  segment  a little  shorter  than  following 
three  segments  combined;  abdominal  terga  with  defi- 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  5 


nite  metallic  bluish  reflections;  punctures  of  scutellum 
mostly  more  separated,  with  many  interspaces  of  more 

than  a puncture  diameter lanosa  Cresson 

10a.  Median  line  of  clypeus  broad,  smooth,  shiny,  and  ini- 
punctate;  punctures  of  disc  mostly  well  separated;  hairs 
at  side  of  dorsal  face  of  first  tergum  short,  light  brown 
anteriorly,  becoming  short  and  fuscous  toward  distal 

margin  angustifrons  Snelling 

b.  Median  line  of  clypeus  narrow,  roughened  and  dull; 
punctures  of  disc  mostly  subcontiguous;  some  hairs  of 
dorsal  face  of  first  tergum  conspicuously  long,  erect, 

plumose,  and  pale  whitish  harbisoni  Snelling 

1 la.  Small  species,  8.5-1 3.5  mm  long;  clypeus  with  distinct 
impunctate  median  line,  punctures  elsewhere  separated 
by  less  than  twice  a puncture  diameter  (if  clypeal  punc- 
tures somewhat  sparse,  scape,  femora,  and  tibiae  fer- 
ruginous)   12 

b.  Larger  species,  15.5-18.5  mm  long;  clypeus  polished 
between  scattered  punctures,  usually  separated  by  more 
than  twice  a puncture  diameter,  and  median  impunc- 
tate line  undefined  (scape,  femora,  and  tibiae  dark 

brownish)  caesa/piniae  Cockerell 

12a.  Color  of  mandible,  labrum,  and  clypeus  usually  not  as 
below;  clypeus  polished  and  shiny,  median  impunctate 
line  present  and  usually  well  defined;  tergal  discs  mod- 
erately to  strongly  shiny,  with  or  without  bluish  reflec- 
tions; scape,  femora,  and  tibiae  brown  or  red  ...  13 
b.  Mandible  (except  apex),  labrum,  and  clypeus  dull  or- 
ange-ferruginous; clypeus  usually  contiguously  punc- 
tate, dull  and  without  median  impunctate  line,  but  may 
be  moderately  shiny  and  with  narrow  impunctate  line; 
tergal  discs  dull  and  closely  tessellate,  with  dull  bluish 

reflections;  scape,  femora,  and  tibiae  brown  

ferrisi  Cockerell 

1 3a.  Basal  face  of  first  tergum  with  abundant  long,  plumose, 
white  hairs,  some  of  which  extend  onto  dorsal  face  in 

middle;  terga  with  definite  bluish  reflections 14 

b.  Basal  face  of  first  tergum  with  sparse  plumose,  white 
hairs,  those  in  middle  shorter  and  not  extending  to 
dorsal  face  in  middle;  terga  black,  without  bluish  re- 
flections   cockerel/i  W.  Fox 

14a.  Mandible,  labrum,  clypeus,  scape,  femora,  and  tibiae 
mostly  or  entirely  bright  ferruginous;  apical  margins  of 
second  and  third  terga  with  short  bands  of  appressed 
pale  hairs;  first  flagellar  segment  shorter  than  following 

three  combined  rhodopus  Cockerell 

b.  Mandible,  scape,  and  legs  brown,  labrum  and  most  of 
clypeus  yellowish;  second  and  third  terga  with  all  hairs 
blackish;  first  flagellar  segment  longer  than  following 

three  combined  atripes  Mocsary 

1 5a.  Clypeus  wholly  black;  pubescence  of  mesepisternum 


wholly  blackish  16 

b.  Clypeus  largely  or  entirely  whitish,  yellowish  or  red- 
dish; pubescence  of  mesepisternum  pale  or  dark  . . 18 

1 6a.  Pubescence  of  thoracic  dorsum  pale  17 

b.  Pubescence  of  thoracic  dorsum  blackish  


aterrima  F.  Smith 

17a.  Pubescence  of  pronotal  lobe,  and  usually  of  entire  lat- 


eral pronotal  area,  pale;  first  flagellar  segment  2.6-3. 1 

times  longer  than  second  zacateca  Snelling 

b.  Pubescence  of  entire  lateral  pronotal  area,  including 
lobe,  blackish;  first  flagellar  segment  3. 8-3. 9 times  longer 

than  second mexicana  F.  Smith 

1 8a.  Pubescence  of  head  and  thorax  at  least  partially  pale 

19 

b.  Pubescence  of  head  and  thorax  blackish  

nigrocaeru/ea  F.  Smith 

1 9a.  Metabasitarsus  much  broadened  toward  apex,  about 
twice  longer  than  greatest  width;  metafemur  swollen 

and  about  twice  longer  than  wide  20 

b.  Metabasitarsus  slender  and  parallel-sided,  at  least  3.5 
times  longer  than  wide;  metafemur  usually  about  three 

times  longer  than  wide,  but  may  be  swollen  22 

20a.  Punctures  equally  dense  on  mesoscutum  and  scutellum; 
mesepisternum  finely,  closely  punctate;  clypeus  pale 

yellow 21 

b.  Punctures  of  mesoscutum  much  sparser  than  those  of 
scutellum;  mesepisternum  with  scattered,  obscure,  fine 

punctures;  clypeus  orange-ferruginous  

ectypha  Snelling 

2 1 a.  First  flagellar  segment  longer  than  following  three  com- 
bined; punctures  of  clypeal  disc  mostly  separated  by  a 
puncture  diameter  or  more  ....  angustifrons  Snelling 
b.  First  flagellar  segment  a little  shorter  than  following 
three  combined;  punctures  of  clypeal  disc  mostly  sep- 
arated by  less  than  0.75  times  a puncture  diameter 

harbisoni  Snelling 

22a.  Discs  of  second  to  fourth  tergites  closely  and  sharply 
punctate,  punctures  separated  by  no  more  than  twice 
a puncture  diameter  and  clearly  greater  in  diameter 

than  hairs  arising  from  them  23 

b.  Discs  of  second  to  fourth  tergites  polished  and  shiny 
between  scattered,  obscure  punctures  little  greater  in 

diameter  than  hairs  arising  from  them  

fisheri  Snelling 

23a.  Clypeus  polished  or  not,  punctures  mostly  separated 
by  two  puncture  diameters  or  less;  ocellocular  distance 
no  more  than  1.25  times  diameter  of  anterior  ocellus; 
smaller  species,  head  width  3. 8-5. 3 mm,  almost  always 

less  than  5.0  mm  24 

b.  Clypeus  polished,  nearly  impunctate;  ocellocular  dis- 
tance at  least  1.65  times  diameter  of  anterior  ocellus; 
larger  species,  head  width  5.0-6. 5 mm,  almost  always 

more  than  5.3  mm  caesalpiniae  Cockerell 

24a.  Clypeus  slightly  shiny  to  shiny,  punctures  ofdisc  mostly 
subcontiguous,  usually  with  distinct  impunctate  me- 
dian line,  sometimes  with  median  line  roughened  and 
dull;  abdominal  terga  without  pale  hairs  beyond  first 
segment;  legs  medium  to  dark  brown  and  metatibia 

mostly  dark  pubescent  25 

b.  Clypeus  polished  between  sparse  punctures  and  median 
impunctate  line  very  broad  and  poorly  defined;  discs 
(or  at  least  apical  margins)  of  second  to  sixth  terga  with 
subappressed  (suberect  in  metander),  mostly  simple, 
pale  hairs;  legs  almost  always  ferruginous  and  metatibia 
at  least  largely  whitish  pubescent . . rhodopus  Cockerell 


6 Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


25a.  Face  narrow,  eye  length  at  least  1 .56  (and  usually  more 
than  1.60)  times  interocular  distance  at  level  of  anten- 
nal sockets;  ocellocular  distance  no  more  than,  and 
usually  less  than,  diameter  of  anterior  ocellus  ...  26 
b.  Face  broad,  eye  length  no  more  than  1.51  (and  usually 
less  than  1.45)  times  interocular  distance  at  level  of 
antennal  sockets;  ocellocular  distance  a little  greater 

than  diameter  of  anterior  ocellus  27 

26a.  Paraocular  area,  mandible  (mostly)  and  underside  of 
scape  (usually)  yellow;  clypeus  closely  punctate  and  me- 
dian impunctate  line  narrow;  abdominal  terga  with 

bluish  reflections  lanosa  Cresson 

b.  Paraocular  area  and  underside  of  scape  dark,  mandible 
ferruginous;  clypeus  with  median  impunctate  line  broad 
and  punctures  mostly  separated  by  about  a puncture 
diameter;  abdominal  terga  black,  without  bluish  re- 
flections   cockerelli  W.  Fox 

27a.  Median  area  of  clypeus  sharply  roughened  and  dull, 
contrasting  to  shiny,  closely  punctate  areas  on  either 
side;  discs  of  abdominal  terga  two  to  five  sharply  tes- 

sellate  and  slightly  shiny  ferrisi  Cockerell 

b.  Clypeal  disc  shiny,  with  distinct  smooth,  impunctate 
median  line;  discs  of  abdominal  terga  two  to  five  shiny, 
not  obviously  tessellate  atripes  Mocsary 

Centris  ( Paracentris ) angustifrons  Snelling 

Centris  ( Paracentris ) angustifrons  Snelling,  1966:13-14.  9. 

The  type  locality  for  C.  angustifrons  is  Huachuca  Mountains, 
Arizona  and  this  species  was  based  on  a single  female  col- 
lected nearly  80  years  ago.  A few  additional  specimens  are 
now  available,  including  the  previously  unknown  males.  The 
males  are  very  similar  to  those  of  C.  harbisoni,  especially  in 
the  shape  of  the  metabasitarsus,  but  are  easily  separated  by 
the  characteristics  cited  in  the  key  above. 

The  females  of  C.  angustifrons  and  C.  harbisoni  are  even 
more  similar,  a fact  I had  not  fully  appreciated  when  I pre- 
pared my  1974  key.  The  present  key  brings  the  two  species 
to  the  same  couplet;  the  differences  between  them  are  noted 
there  and  should  present  few  difficulties. 

NEW  RECORDS 

MEXICO,  SONORA:  13,  Aduana,  1 5 Mar.  1 962  (L.A.  Stange, 
UCD);  699,  233,  Rio  Cuchuhaqui,  8 mi.  S Alamos,  1-13  Apr. 
1975  (A.  Brewster;  LACM),  on  Parkinsonia  sp.  (299,  13), 
Fourquieria  sp.  (13),  Cercidium  sp.  (299),  and  Prosopis  sp. 
(299). 

Centris  ( Paracentris ) aterrima  F.  Smith 

Centris  aterrima  F.  Smith,  1854:378.  3. 

Centris  (Paracentris)  aterrima:  Snelling,  1974:7,  8 (key). 

NEW  RECORDS 

UNITED  STATES,  ARIZONA,  Cochise  Co.:  19,  Rustler  Park, 
Chiricahua  Mountains,  4 Aug.  1971  (LACM);  699,  1 mi.  E 
Douglas,  14  Aug.  1969  (J.G.  and  K.C.  Rozen;  AMNH);  19, 
Yaqui  Canyon  area,  5370-5700  ft.  elev.,  Huachuca  Moun- 


tains, 29  Aug.  1972  (R.R.  Snelling;  LACM),  on  Acacia  an- 
gustissima.  PimaCo.:  299,  Arivaca,  1 8 July  and  24  Aug.  1974 
(J.L.  Neff;  LACM).  Santa  Cruz  Co.:  19,  13,  Sycamore  Can- 
yon, near  Ruby,  16-17  Aug.  1961  (J.C.  Bequaert;  LACM); 
13,  Nogales,  24  Aug.  1939  (R.H.  Crandall;  LACM);  13,  Pat- 
agonia, 24  Aug.  1955  (F.G.  Werner  and  G.D.  Butler;  LACM); 
19,  White  Rock  Campground,  0.5  mi.  S Pena  Blanca,  9 Aug. 
1972  (D.C.  Frack;  LACM);  19,  Pena  Blanca,  27  July  1972 
(D.C.  Frack;  LACM).  MEXICO,  CHIAPAS:  19,  333,  12  mi. 
W Ocozocoautla,  26  July  1953  (E.E.  Gilbert  and  C.D. 
MacNeill;  UCB).  OAXACA:  13,  4 mi.  N Pochutla,  150  m 
elev.,  1 1 Oct.  1975  (J.L.  Neff;  LACM),  on  “Mint  71  15”;  399, 
4 mi.  W Zanatepec,  200  m elev.,  16  Sept.  1975  (J.L.  Neff; 
LACM.  NEFF),  on  Krameria  revoluta.  SINALOA:  19,  30  mi. 
E Villa  Union,  570  m elev.,  20  Mar.  1980  (J.L.  Neff;  NEFF). 

Centris  ( Paracentris ) atripes  Mocsary 

Centris  atripes  Mocsary,  1899:254:  3. 

Centris  limbata  Friese,  1899:44.  9.  NEW  SYNONYMY. 
Centris  atriventris  W.  Fox,  1899:68.  9 3.  Preoccupied. 
Centris  Foxi  Friese,  1900b:350.  New  name  for  C.  atriventris 
W.  Fox. 

Centris  (Paracentris)  atripes:  Snelling,  1974:8,  9-10  (key, 
distr.,  var.). 

Friese  described  C.  limbata  from  a single  female  collected 
by  G.  Birkmann  at  Fedor,  Lee  County,  Texas.  The  most 
distinctive  feature  of  this  species  was  the  presence  of  distinct 
bands  of  pale  hairs  on  the  metasomal  terga.  Since  no  spec- 
imens resembling  the  description  have  been  subsequently 
collected.  C.  limbata  remained  an  enigma. 

At  my  request,  R.W.  Brooks  examined  the  type  of  C.  lim- 
bata in  the  Berlin  Museum  and  he  provided  several  photo- 
graphs and  sketches.  A photograph  of  the  lateral  view  of  the 
type  specimen  clearly  shows  that  the  abdomen  is  glued  to 
the  thorax.  The  abdomen  is  distinctly  fasciate  in  dorsal  view 
and  a prominent,  apically  truncate  pygidial  plate,  without  a 
secondary  plate,  is  present.  In  fact,  both  the  abdominal  bands 
and  the  pygidial  plate  are  typical,  not  of  Centris,  but  of  a 
Me/issodes  male!  The  head  and  thorax  of  the  type  are  just 
as  clearly  those  of  a C.  atripes  female.  Since  the  species  was 
described  as  a Centris,  1 consider  these  parts  to  be  the  true 
type;  the  abdomen  is  extraneous.  This  restriction  negates  any 
possible  nomenclatural  confusion  within  Me/issodes. 

NEW  RECORDS 

COSTA  RICA,  GUANACASTE:  19,  8 km  NW  Liberia,  9 
Feb.  1975  (G.R.  Frankie;  TAMU),  on  Cassia  biflora,  0900. 
GUATEMALA:  19,  8 mi.  NE  El  Progreso,  8 July  1965  (A. 
Raske  and  C.  Slobodchikoff;  UCB);  13,  Jicara,  8 May  1931 
(J.  Bequaert;  AMNH).  UNITED  STATES,  OKLAHOMA: 
19,  Norman,  Cleveland  Co.,  “8/4  1949”  (W.T.  Nailon; 
UKAN). 

Centris  ( Paracentris ) cockerelli  W.  Fox 

Centris  lanosa:  W.  Fox,  1899:69;  Cockerell,  1906:97;  Lutz 
and  Cockerell,  1 920:556;  Timberlake,  1940: 1 38;  Snelling, 
1956:7  (in  part,  misidentifications). 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  7 


Centris  cockerelli  W.  Fox,  1899:68.  9. 

Centris  cockerelli  resoluta  Cockerell,  1923:76-77.  9 <3.  NEW 
SYNONYMY. 

Centris  lanosa  lanosa:  Snelling,  1966:6  (misidentification). 
Centris  lanosa  resoluta:  Snelling,  1966:6. 

Centris  (Paracentris)  cockerelli  cockerelli:  Snelling,  1974: 1 0- 
1 1. 

Centris  (Paracentris)  cockerelli  resoluta:  Snelling,  1974: 1 1 . 

For  complete  literature  citations  of  this  common  species,  see 
Snelling  (1974)  under  C.  cockerelli  and  C.  c.  resoluta.  This 
is  the  species  long  improperly  called  C.  lanosa.  Although  I 
have  attempted  to  maintain  recognition  of  C.  c.  resoluta.  1 
no  longer  believe  this  to  be  correct.  In  the  females  there  are 
two  principal  phenotypes:  an  eastern  form  with  whitish  to 
yellowish  clypeus  and  a western  form  with  a reddish  yellow 
clypeus.  There  are  no  apparent  differences  to  be  seen  in  the 
males  associated  with  these  females.  The  two  female  forms 
intergrade  continuously  from  western  Texas  and  Tamaulipas 
through  New  Mexico  and  Chihuahua.  In  my  opinion  these 
variants  are  the  end-points  of  a cline  of  variation  and  are 
not  worthy  of  formal  separation. 

Centris  ( Paracentris ) ferrisi  Cockerell 

Centris  atripes  subsp.  ferrisi  Cockerell,  1924:49.  “9”  = <3! 

This  was  described  from  two  specimens  from  La  Paz.  Baja 
California  Sur,  Mexico;  although  Cockerell  stated  the  spec- 
imens to  be  females,  both  are  males.  The  types  are  in  the 
CAS.  In  my  key  (1974),  the  males  will  run  to  C.  atripes. 
Males  differ  from  those  of  C.  atripes.  and  other  species  of 
similar  appearance,  in  that  the  integument  of  the  abdominal 
tergites  is  dull  and  sharply  roughened  between  minute,  ob- 
scure punctures;  the  clypeus,  especially  basad.  is  distinctly 
roughened  between  well-separated  punctures.  The  females, 
too,  will  key  to  C.  atripes  and,  as  in  the  male,  have  the  tergal 
sculpture  distinct,  dulling  the  surface;  the  facial  marks  are 
dull  ferruginous  rather  than  yellow  as  in  C.  atripes',  in  C. 
ferrisi  the  first  flagellar  segment  is  as  long  as  the  following 
three  combined,  shorter  in  C.  atripes.  The  male  clypeus  is 
usually  ferruginous,  but  is  yellow  in  some  specimens. 

In  addition  to  the  types  of  C.  ferrisi  I have  seen  the  fol- 
lowing, all  from  MEXICO,  BAJA  CALIFORNIA  SUR:  1 <3, 
Cabo  San  Lucas,  8-14  Sept.  1978  (J.P.  and  K.E.  Donahue; 
LACM);  19,  1<3,  Canon  de  la  Zorra,  260  m elev.,  1 1 km  W 
Santiago,  4-5  Sept.  1977  (R.R.  Snelling;  LACM),  on  Par- 
kinsonia  aculeata\  299,  4 km  N Los  Barriles,  10  m elev.,  4 
Sept.  1 977  (R.R.  Snelling;  LACM);  19,  3.7  mi.  W La  Burrera, 
1400  ft.  elev.,  7-8  Oct.  1975  (R.R.  Snelling;  LACM),  on 
Antigonon  leptopus ; 1 <3,  2.5  mi.  SE  La  Huerta,  2200  ft.  elev., 
8-9  Oct.  1968  (E.L.  Sleeper  and  F.J.  Moore;  LACM);  19,  2 
mi.  S La  Paz,  6 Aug.  1966  (J.A.  Chemsak;  UCB),  on  P. 
aculeata ; 13,  7 mi.  SW  La  Paz  (J.A.  Chemsak;  UCB),  on 
Wislizenia  refract  a var.  mamillata\  19,  23  km  W La  Paz, 
24-27  Apr.  1975  (E.M.  and  J.L.  Fisher;  LACM);  19,  68  km 
S Loreto,  Km  76  on  Hwy  1,  29  July  1977  (D.  Weismann 
and  C.  Mullinex;  CAS);  13,  La  Paz,  15  Sept.  1983  (R.R. 
Snelling;  LACM),  on  A.  leptopus',  19,  25  mi.  W La  Paz,  30 


Aug.  1959  (E.W.  Radford  and  F.G.  Werner;  CAS);  19,  3 mi. 
E San  Pedro,  15  Sept.  1983  (R.R.  Snelling;  LACM),  on  A. 
leptopus\  13,  52  mi.  NW  La  Paz,  15  Mar.  1980  (J.L.  Neff; 
NEFF),  on  Cercidium  penmsulare:  19,  La  Laguna  (Sierra  de 
la  Laguna),  1829  m elev.,  23  Oct.  1977  (D.E.  and  W.R. 
Breedlove;  CAS);  13,  11  mi.  NE  Todos  Santos,  16  Sept.  1983 
(R.R.  Snelling;  LACM),  on  A.  leptopus ; 19,  25  km  E Todos 
Santos,  near  La  Burrera,  1829  m elev.,  21  Oct.  1977  (D.E. 
and  W.R.  Breedlove;  CAS);  13,  Miraflores,  17  Sept.  1983 
(R.R.  Snelling;  LACM),  on  A.  leptopus ; 19,  1 mi.  S Agua 
Caliente,  17  Sept.  1983  (R.R.  Snelling;  LACM),  on  A.  lep- 
topus', 299,  5 mi.  W San  Ignacio,  13  Sept.  1983  (R.R.  Snelling; 
LACM),  on  Tephrosta  tenella ; 299,  same,  except  19  Sept. 
1983. 

Centris  ( Paracentris ) fisheri  Snelling 

Centris  ( Paracentris ) fisheri  Snelling,  1974:12.  3. 

This  was  described  from  two  males  collected  near  San  Ig- 
nacio, Baja  California  Sur,  Mexico.  The  female  will  go  to, 
and  closely  resembles,  C.  harbisoni  Snelling  in  my  key  (1974) 
to  species  of  Paracentris.  but  differs  immediately  in  that  the 
abdominal  terga  are  polished  and  very  sparsely  punctate;  in 
C.  fisheri  the  first  flagellar  segment  is  shorter  than  the  scape 
and  shorter  than  the  combined  lengths  of  the  second  to  fourth 
flagellar  segments;  the  first  flagellar  segment  is  longer  in  C. 
harbisoni. 

NEW  RECORDS 

MEXICO,  BAJA  CALIFORNIA  SUR:  1 199,  533,  5 mi.  W 
San  Ignacio,  13  Sept.  1983  (R.R.  Snelling;  LACM),  on  Te- 
phrosia  tenella',  599,  1033,  same,  except  19  Sept.  1983;  19, 
19  km  NW  Mulege,  8 Sept.  1977  (R.R.  Snelling;  LACM),  on 
Hojfmanseggia  sp.;  13,  35  mi.  N Loreto,  5 Oct.  1975  (R.R. 
Snelling;  LACM),  on  Wislizenia  refract a\  533,  9.6  mi.  N Lore- 
to, 14  Sept.  1983  (R.R. Snelling;  LACM),  on  Antigonon  lep- 
topus', 19,  Estacion  Microondas  “Ligui,”  48  km  S Loreto,  425 
melev.,  14  Sept.  1 983  (R.R.  Snelling;  LACM),  on  A.  leptopus: 
233,  68  km  S Loreto,  Km  76  on  Hwy  1,  29  July  1977  (D. 
Weismann  and  C.  Mullinex;  CAS). 

Centris  ( Paracentris ) harbisoni  Snelling 

Centris  (Paracentris)  harbisoni  Snelling,  1974: 14-16.  9 3. 

NEW  RECORDS 

MEXICO,  BAJA  CALIFORNIA  SUR:  19,  San  Jose  del  Cabo, 
11-16  Sept.  1967  (J.  Chemsak,  A.  and  M.  Michelbacher; 
UCB);  13,  52  mi.  NW  La  Paz,  1 5 Mar.  1980  (J.L.  Neff;  NEFF) 
on  Cercidium  peninsulare',  499,  106  km  N La  Paz,  18  Mar. 
1980  (J.L.  Neff;  NEFF),  on  Krameria  parvifolia. 

Centris  (Paracentris)  laevibullata  Snelling 

Centris  (Paracentris)  laevibullata  Snelling,  1966:17-18.  9; 
Snelling,  1974:7  (key). 

This  species  was  described  from  a female  from  Orizaba,  Vera 


8 Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


Cruz,  Mexico  (type  locality)  and  another  collected  14  mi. 
NW  Zitacuaro,  Michoacan,  Mexico.  The  male  is  unknown. 

NEW  RECORDS 

MEXICO,  DUR.ANGO:  19,  Durango,  13  Aug.  1962  (A.E. 
Michelbacher;  UCB).  JALISCO:  19,  3.5  mi.  E [Rio]  Mag- 
dalena, 5 Sept.  1965  (A.R.  Gillogly;  LACM);  19,  25  mi.  W 
Guadalajara,  4700  ft.  elev.,  29  Sept.  1957  (H.A.  Scullen; 
ORSU);  19,  13  mi.  NW  Lagos  de  Moreno,  3 Sept.  1975  (J.L. 
Neff;  LACM).  MICHOACAN:  19,  7.7  km  NE  Patzcuaro, 
2088  m elev.,  23  Sept.  1 976  (C.D.  George  and  R.R.  Snelling; 
LACM).  NUEVO  LEON:  19,  18  mi.  W Linares,  2700  ft. 
elev.,  26  Sept.  1975  (J.A.  Powell,  J.  Chemsak,  and  T.  Fried- 
lander;  UCB).  SAN  LUIS  POTOSI:  19,  52  mi.  S.  Tamazun- 
chali,  5700  ft.  elev.,  7 Oct.  1957  (H.A.  Scullen;  ORSU). 

Centris  ( Paracentris ) lanosa  Cresson 

Centris  lanosa  Cresson,  1872:284.  3. 

Centris  subhyalina  W.  Fox,  1899:69.  9. 

Centris  birkmanii  Friese,  1899:44.  <5  9. 

Centris  (Paracentris)  lanosa:  Snelling,  1974:8,  16-17  (key, 
syn.). 

NEW  RECORDS 

UNITED  STATES,  FLORIDA:  19,  Austin  Carey,  Alachua 
Co.,  22-24  May  1975  (G.B.  Fairchild;  DPIF);  19,  Gainesville, 
Alachua  Co.  (DPIF);  19,  Trenton,  Gilchrist  Co.,  14  Apr.  1925 
(D.M.  Bates;  DPIF).  KANSAS:  19,  3 mi.  S Sawyer,  Pratt  Co., 
16  June  1962  (C.D.  Michener  and  party;  UKAN),  on  Amor- 
pha  canescens.  OKLAHOMA:  19,  near  Ardmore,  Carter  Co., 
3 June  1961  (Umv.  Kans.  Mex.  Exped.;  UKAN);  19,  Cleve- 
land Co.,  31  May  1951  (H.L.  Parker;  UKAN);  19,  Ft.  Sill, 
Comanche  Co.,  24  June  1974(T.E.  Rogers;  LACM).  TEXAS: 
599,  Bastrop,  Bastrop  Co.,  2 May  1969  (Brothers,  Krueger, 
Michener;  UKAN). 

Centris  ( Paracentris ) mexicana  F.  Smith 

Centris  mexicana  F.  Smith,  1854:378.  “9”  = <5! 

Centris  ( Paracentris ) mexicana:  Snelling,  1974:7,  8,  17-18 
(key,  tax.,  distr.). 

NEW  RECORDS 

MEXICO,  OAXACA:  499,  233,  Tamazulapam,  6200  ft.  elev., 
2 Sept.  1965  (S.J.  Arnold;  UCB),  on  Salvia  sp.;  13,  Monte 
Alban  ruins,  3 Aug.  1964  (H.V.  Daly;  UCB).  VERA  CRUZ: 
19,  2 66,  Cotaxtla  Exp.  Sta.,  Cotaxtla,  9 Aug.  1962  (D.H. 
Janzen;  UCB). 

Centris  ( Paracentris ) nigrocaerulea  F.  Smith 

Centris  nigro-caerulea  F.  Smith,  1874:369.  9 6. 

Centris  clypeata  Friese,  1899:41.  9 <3.  Preoccupied.  NEW 
SYNONYMY. 


Centris  (Paracentris)  anthracina  Snelling,  1966:14-17.  9 6. 
NEW  SYNONYMY. 

DISCUSSION 

I have  examined  the  female  and  male  type  specimens  of  C. 
nigro-caerulea  and  they  are  conspecific  with  C.  clypeata  and 
C.  anthracina.  Smith’s  types  are  from  an  unspecified  locality 
in  Mexico;  the  female  is  herewith  designated  lectotype  and 
the  male  as  paralectotype.  Both  are  in  the  collections  of  the 
British  Museum  (Natural  History). 

NEW  RECORDS 

MEXICO,  CHIAPAS:  333,  733,  3 mi.  W Navenchuac,  1-2 
Apr.  1953  (R.C.  Bechtel  and  E.  I.  Schlinger;  UCB);  19,  be- 
tween Tuxtla  [Gutierrez]  and  Chilapa,  23  Jan.  1 974  (S.  Buch- 
mann;  LACM),  on  Cassia  sp.  JALISCO:  333,  “env.  de  Gua- 
dalajara,” 1903  and  May  1913  (L.  Diguet;  MNHN). 
MORELOS:  19,  Cuernavaca,  6000  ft.  elev.,  7 Feb.  1954 
(R.R.  Dreisbach;  LIKAN).  OAXACA:  13,  Oaxaca,  12  Oct. 
1963  (A.E.  and  M.M.  Michelbacher;  UCB);  499,  20  mi.  S 
Taxco,  6 Feb.  1954  (R.R.  Dreisbach;  UKAN).  SONOR. 4: 
13,  Aduana  (near  Alamos),  15  Mar.  1962  (F.D.  Parker;  UCD); 
13,  Rio  Cuchuhaqui,  8 mi.  S Alamos,  1-13  Apr.  1975  (A. 
Brewster;  LACM),  on  Parkinsonia  sp.  GUATEMALA:  399, 
San  Miguel  Duenas,  5200  ft.  elev.,  1-7  Dec.  1975  (S.W.T. 
Batra;  LACM);  19,  Ciudad  de  Guatemala,  Dec.  1911  (W.M. 
Wheeler;  MCZ);  19,  Universidad  del  Valle,  Ciudad  de  Gua- 
temala, 25  Nov.  1975  (S.W.T.  Batra;  LACM).  PANAMA. 
CHIRIQUI:  2 33,  Potrero  Grande,  28  Jan.  1981  (D.W.  In- 
ouye;  ROUB). 

Subgenus  Acritocentris  Snelling,  1974 

Centris  subg.  Acritocentris  Snelling,  1974:36.  Type-species: 
Centris  (Melanocentris)  ruthannae  Snelling,  1966;  original 
designation. 

In  the  years  subsequent  to  the  description  of  this  subgenus 
and  my  treatment  of  its  component  species,  I have  been  able 
to  examine  many  more  specimens  than  were  then  available. 
As  a result,  I can  now  provide  a more  adequate  account  of 
these  species.  The  key  given  then  (Snelling,  1974)  will  not 
always  work  for  the  females  and  should  be  replaced  by  the 
new  one  given  below. 

KEY  TO  SPECIES  OF  ACRITOCENTRIS 

1 a.  Female,  antenna  1 2-segmented  and  basitibial  plate  pres- 
ent   2 

b.  Male,  antenna  13-segmented  and  basitibial  plate  absent 

4 

2a.  Pubescence  of  thoracic  dorsum  whitish  3 

b.  All  pubescence  blackish  ruthannae  Snelling 

3a.  First  abdominal  tergite  nearly  im punctate  toward  apical 
margin,  especially  in  middle;  median  area  of  clypeus  dull 
and  roughened  but  not  rugose,  laterad  with  rounded, 
separated  punctures;  basal  shiny  area  of  clypeus  broad- 
ened in  middle  a/biceps  Friese 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  9 


b.  First  abdominal  tergite  closely,  though  minutely,  punc- 
tate to  apical  margin;  median  area  of  clypeus  rugose  and 
dull,  laterad  with  coarse,  close,  elongate  punctures;  basal 
shiny  area  narrow  across  entire  clypeal  base  


agameta  Snelling 

4a.  Thoracic  dorsum  dark  pubescent  5 

b.  Thoracic  dorsum  pale  pubescent  6 

5a.  Labrum,  clypeus,  paraocular  area,  and  supraclypeal  area 

all  yellow-maculate  ruthannae  Snelling 

b.  Entire  face  black  satana,  new  species 


6a.  Face  marks  whitish,  absent  from  paraocular  and  supra- 
clypeal areas,  clypeus  broadly  blackish  along  lateral  and 
basal  margin,  disc  smooth  and  shiny  . . . a/biceps  Friese 
b.  Face  marks  yellowish,  usually  present  on  paraocular  and 
supraclypeal  areas,  lateral  and  basal  margins  of  clypeus 
narrowly,  or  not  at  all.  blackish,  disc  dull  and  roughened 
agameta  Snelling 

Centris  ( Acritocentris ) albiceps  Friese 

Centris  mexicana  var.  albiceps  Friese,  1899:289.  3. 

Centris  (Melanocentris)  strawi  Snelling,  1 966:27-28.  <5.  NEW 
SYNONYMY. 

Centris  ( Acritocentris ) strawi  Snelling,  1974:37,  38  (in  part) 
(key,  distr.). 

1 suggested  (1974)  that  C.  mexicana  var.  albiceps  might  be 
an  older  name  for  C.  strawi.  Friese  based  this  name  on  a 
specimen  from  an  unknown  locality  in  Mexico.  The  type  is 
in  the  Berlin  Museum  and  was  examined  at  my  request  by 
R.W.  Brooks  who  confirmed  the  above  synonymy.  The  type 
is  not  so  marked,  and  bears  two  labels:  “116  59/Mexico” 
and  “ Centrist  mexicana/ v . albiceps/Friese  1898.” 

The  females  which  in  1974  1 believed  to  belong  to  C. 
albiceps  (as  C.  strawi)  are  now  known  to  be  those  of  C. 
agameta.  The  true  females  of  this  species  are  very  similar  to 
those  of  C.  agameta.  but  the  disc  of  the  clypeus,  although 
dull  and  roughened,  lacks  the  characteristic  irregular  rugulae 
present  in  C.  agameta.  The  first  two  abdominal  terga  are 
much  more  weakly  punctate  toward  the  margins  in  C.  al- 
biceps. and  in  C.  albiceps  females  the  labrum  and  posterior 
pronotal  lobes  are  without  pale  hairs. 

NEW  RECORDS 

MEXICO,  CHIAPAS:  329,  9<5<5,  Municipio  Chiapo  de  Corzo, 
El  Chorreadero,  753  m elev.,  1 Nov.  1976  (D.E.  and  J.A. 
Breedlove;  CAS);  12,  Municipio  Motozintla,  betw.  Motozin- 
tla  and  Mazapa,  1219  m elev.,  5 Oct.  1976  (D.E.  and  J.A. 
Breedlove;  CAS).  HIDALGO:  229,  Zimapan,  6400  ft.  elev., 
8 Oct.  1957  (H.A.  Scullen;  ORSU).  MICHOACAN:  229,  Lake 
Patzcuaro,  6800  ft.  elev.,  2 1 Sept.  1 957  (H.A.  Scullen;  ORSU); 
13,  10  mi.  N Morelia,  5900  ft.  elev.,  28  July  1962  (Univ. 
Kans.  Mex.  Exped.;  UKAN),  on  Leguminoseae;  12,  233,  Ca- 
rapan,  1 Sept.  1962  (D.H.  Janzen;  UCB).  NUEVO  LEON: 
12,  30  mi.  N Linares,  1500  ft.  elev.,  11  Oct.  1957  (H.A. 
Scullen;  ORSU).  OAXACA:  13,  22  mi.  SE  Oaxaca,  5700  ft. 
elev.,  2 Sept.  1957  (H.A.  Scullen;  ORSU);  12,  5 mi.  NW 
Totolapan,  3800  ft.  elev.,  6 July  1953  (Univ.  Kans.  Mex. 


Exped.;  UKAN)  on  Lonchocarpus\  522,  4 mi.  N Totolapan, 
1849  m elev.,  15  Sept.  1975  (J.L.  Neff;  LACM,  NEFF),  on 
“ Cassia  7064”;  12,  2 mi.  S Totolapan,  15  Sept.  1975  (J.L. 
Neff;  LACM),  on  “ Cassia  7064”;  13,  near  Las  Margaritas, 
1400  m elev.,  15  Sept.  1975  (J.L.  Neff;  LACM).  PUEBLA: 
13,  3 mi.  NW  Petlalcingo,  4600  ft.  elev.,  5 Sept.  1972  (Byers 
and  Thornhill;  UKAN);  12,  22  km  NW  Izucar  de  Matamoros, 
1158m  elev.,  2 1 Sept.  1 976  (C.D.  George  and  R.R.  Snelling; 
LACM),  on  Cassia  laevigata.  QUERETARO:  12,  10  mi.  S 
Jet.  Hwy  55  and  45,  30  Aug.  1963  (Scullen  and  Bollinger; 
ORSU).  SAN  LUIS  POTOSI:  12,  9 mi.  E Ciudad  [del]  Maiz, 
3975  ft.  elev.,  23  July  1962  (Univ.  Kans.  Mex.  Exped.; 
UKAN);  12,  5 mi.  E Ciudad  [del]  Maiz,  4700  ft.  elev.,  22 
Aug.  1954  (Univ.  Kans.  Mex.  Exped.;  UKAN);  12,  1 133,  6 
mi.  W Guadalcazar,  2 Sept.  1975  (J.L.  Neff;  LACM,  NEFF), 
on  “mint  7031.”  TAMAULIPAS:  3622,  38  mi.  N El  Mante, 
1050  ft.  elev.,  1 1 Oct.  1957  (H.A.  Scullen;  ORSU);  1622,  9, 
12,  and  24  mi.  S [Ciudad]  Victoria,  11  Oct.  1957  (H.A. 
Scullen;  ORSU);  422,  15  mi.  SW  Ciudad  Victoria,  5000  ft. 
elev.,  19  Sept.  1976  (J.A.  Chemsak  et  al.;  UCB);  2022,  35 
mi.  S Villagran.  15  Sept.  1977  (Chemsak  and  Michelbachers; 
UCB),  on  Solanum\  12,  7 mi.  S Villagran,  26  Sept.  1975  (J. 
Powell  et  al.;  UCB);  12,  1 5 mi.  NE  Juamava,  2500  ft.  elev., 
19  Sept.  1976  (J.  Chemsak  et  al.;  UCB).  VEIL 4 CRUZ:  13, 
3 mi.  E Acultzingo,  1 600  m elev.,  2 1 Aug.  1 977  (E.I.  Schlin- 
ger;  UCB);  12,  233,  E Citlaltepetl,  6000  ft.  elev.,  25  June  1964 
(L.W.  Swan;  CAS). 

Centris  ( Acritocentris ) agameta  Snelling 

Centris  ( Acritocentris ) agameta  Snelling.  1974:37-38.  3. 

NEW  RECORDS 

MEXICO,  GUERRERO:  12,  Acapulco,  6 Aug.  1954  (Univ. 
Kans.  Mex.  Exped.;  UKAN).  JALISCO:  322,  13,  Hwy  15, 
0.2  mi.  from  Nayarit  State  line,  11  Aug.  1963  (D.  Byers; 
UKAN);  322,  6 mi.  NE  El  Rincon,  3 Aug.  1971  (E.M.  Fisher; 
LACM);  12,  Puente  Grande,  5000  ft.  elev.,  20  Aug.  1954 
(Univ.  Kans.  Mex.  Exped.;  UKAN);  12,  Barra  de  Navidad, 
6 Sept.  1966  (E.M.  Fisher;  UKAN);  13,  3 mi.  NW  Tequila, 
4000  ft.  elev.,  15  July  1953  (Univ.  Kans.  Mex.  Exped.; 
LACM),  on  Vitex  pyramidata:  13,  8 km  W Tequila,  18  July 
1951  (P.D.  Hurd;  UCB);  13,  7 km  N Tequila,  6 Sept.  1975 
(J.L.  Neff;  LACM);  13,  14  mi.  NW  [Rio]  Magdalena,  3500 
ft.  elev.,  19  July  1953  (Univ.  Kans.  Mex.  Exped.;  UKAN), 
on  Vitex pyramidata.  MORELOS:  12,  14  mi.  S Cuernavaca, 
3 Aug.  1954  (Univ.  Kans.  Mex.  Exped.;  UKAN);  12,  12  mi. 
E Cuernavaca,  4300  ft.  elev.,  14  Aug.  1954  (Univ.  Kans. 
Mex.  Exped.;  UKAN);  12,  8.7  mi.  NE  Yautepec,  4400  ft. 
elev.,  15  Aug.  1963  (Ordway  and  Roberts;  UKAN);  on  Cu- 
phea  sp.;  13,  6.7  mi.  S Yautepec,  29  July  1963  (Naumann 
and  Willis;  UKAN).  NAYARIT:  322,  Ixtlan  del  Rio,  5 Aug. 
1963  (P.  Fonda-Bonardi;  LACM).  OAXACA:  322,  13,  Salina 
Cruz,  7 Sept.  1965  (D.H.  Janzen;  UKAN);  433,  20  mi.  E El 
Camaron,  7 Aug.  1956  (J.W.  MacSwain;  UCB).  PUEBLA: 
12,  13,  7 mi.  N Izucar  de  Matamoros,  4450  ft.  elev.,  19  Aug. 
1962  (Univ.  Kans.  Mex.  Exped.;  UKAN),  on  Cuphea  sp. 
SAN  LUIS  POTOSI:  13,  9 mi.  E Ciudad  [del]  Maiz,  3975 


10  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


ft.  elev.,  23  July  1962  (Univ.  Kans.  Mex.  Exped.;  UKAN). 
TAMAULIPAS:  433,  40  km  N Soto  la  Marina,  5 Sept.  1975 
(E.M.  and  J.L.  Fisher;  LACM).  SINALOA:  12,  733,  31  km 
N Mazatlan,  76  m elev.,  29  Sept.  1976  (C.D.  George  and 
R.R.  Snelling;  LACM),  on  Antigonon  leptopus. 

Centris  ( Acritocentris ) ruthannae  Snelling 

Centris  ( Melanocentris ) ruthannae  Snelling,  1 966:28-30.  <3  2. 
Centris  ( Acritocentris ) ruthannae:  Snelling,  1 974:37, 40  (distr.). 

This  species  has  been  previously  known  only  from  southern 
Arizona. 

NEW  RECORDS 

MEXICO,  CHIHUAHUA:  12,  Temoris,  13  Sept.  1970  (T.A. 
Sears  et  al.;  UCD).  SINALOA:  12,  2.5  mi.  W Concordia,  25 
Sept.  1977  (J.A.  Chemsak,  A.  and  M.  Michelbacher;  UCB). 
SONOR.A:  12,  Bahia  San  Carlos,  20  July  1965  (D.S.  Verity; 
LACM);  12,  20  km  N Guaymas  (Km  141),  16  Aug.  1979 
(E.M.  Fisher;  LACM);  12,  13,  Navajoa,  12  Sept.  1964  (A.E. 
and  M.M.  Michelbacher;  UCB),  on  morning  glory,  0750- 
0800;  222,  233,  Cerro  Masiaco,  ca.  1 1 road  mi.  SSE  Baca- 
bachi,  ca.  750  ft.  elev.,  29  Aug.  1 976  (J.P.  and  K.E.  Donahue; 
LACM);  13,  37  km  SE  Bacabachi  (Km  90),  14  Aug.  1976 
(E.M.  Fisher;  LACM),  on  Caesa/pinia  sp. 

Centris  ( Acritocentris ) satana,  new  species 

Figures  1-4 

DIAGNOSIS 

Male  unique  within  Acritocentris  by  the  entirely  blackish 
integument  and  pubescence;  female  unknown. 

DESCRIPTION 

HOLOTYPE  MALE.  Measurements  (mm).  Head  width 
6.15  (5.33-5.74);  head  length  4.56  (4.00-4.10);  wing  length 
13.5  (13.0-13.5);  total  length  1 8.0  ( 1 5.0-1 7.0). 

Head.  1.35  (1.33-1.40)  times  broader  than  long;  occipital 
margin,  in  frontal  view,  very  weakly  convex  and  barely  raised 
above  level  of  tops  of  eyes;  ocelli  well  below  occipital  margin 
in  frontal  view;  inner  orbits  weakly  convergent  above,  upper 
frontal  width  0.91  (0.85-0.87)  times  lower  frontal  width. 
Mandible  slender,  tridentate.  Labrum  about  two-thirds  as 
long  as  broad,  apical  margin  narrowly  rounded;  disc  shiny, 
moderately  rugosopunctate  to  contiguously  punctate.  Disc  of 
clypeus  obtusely  raised  along  midline  over  basal  one-half, 
integument  weakly  to  moderately  shiny,  surface  with  dense, 
moderate  punctures  or  with  very  irregular,  fine,  anastomos- 
ing rugules;  punctures  on  side  of  clypeus  subcontiguous,  elon- 
gate. Remainder  of  head  shiny  between  fine  to  moderate, 
dense  to  subcontiguous  punctures  (sparse  mesad  in  ocellocu- 
lar  area),  but  sparse,  minute  and  obscure  over  most  of  gena. 
Interantennal  distance  2.35  (2. 1 3-2.33)  times  antennal  sock- 
et diameter;  antennocular  distance  1.00  (0.87-0.97)  times 
antennal  socket  diameter;  scape  moderately  stout,  2.36  (2.29- 
2.42)  times  longer  than  wide,  scape  length  1 .06  ( 1 .08-1 . 1 4) 


times  length  of  first  flagellar  segment;  first  flagellar  segment 
shorter  than  length  of  following  three  segments  combined, 
4.00  (3.22-4.06)  times  length  of  second;  interocellar  distance 
1 .79  ( 1 .6 1-1 .90)  times  diameter  of  anterior  ocellus;  ocellocu- 
lar  distance  1 .29  (0.94-1 . 1 6)  times  diameter  of  anterior  ocel- 
lus; ocelloccipital  distance  2.21  (1.97-2.09)  times  diameter 
of  anterior  ocellus. 

Thorax.  Mesoscutum  shiny  between  subcontiguous  to 
dense,  moderate  punctures;  scutellum  similar,  but  punctures 
contiguous  to  subcontiguous  and  slightly  elongate.  Mesepi- 
sternum  moderately  shiny,  punctures  dense,  fine  and  oblique. 
Metepisternum  shiny,  punctures  fine  and  sparse  anteriorly, 
becoming  dense  to  subcontiguous  along  posterior  margin. 
Metafemur  stout,  more  than  twice  longer  than  thick;  meta- 
basitarsus  about  three  times  longer  than  broad. 

Abdomen.  Tergal  discs  shiny  between  dense  to  sparse,  fine 
to  moderate  punctures  which  become  sparser  mesad  and  on 
succeeding  segments;  punctures  in  apical  zone  minute.  Py- 
gidial  plate  narrowly  truncate  to  shallowly  notched  at  apex. 

Terminalia.  Seventh  sternite  (Fig.  1)  with  lateral  margins 
of  distal  process  convex,  apical  margin  concave,  with  definite 
mesoapical  row  of  short  hairs.  Eighth  sternite  (Fig.  2)  with 
distal  process  slender,  compressed;  hairs  numerous,  long, 
coarsely  plumose.  Dorsal  process  of  gonocoxite  (Fig.  4)  long, 
slender,  nearly  straight,  slightly  hooked  at  apex;  gonostylus 
slender,  setae  long  and  coarse  along  basal  margin,  shorter 
and  finer  distad. 

Pilosity.  Hairs  uniformly  dark  brown  to  blackish. 

Color.  Blackish  brown,  abdominal  terga  with  strong  bluish 
reflections.  Mandible,  antenna  and  legs  reddish  brown,  fem- 
ora more  conspicuously  reddish.  Wings  dark  brown,  veins 
and  stigma  black. 

TYPE  MATERIAL 

Holotype  male:  Tepoxtlan,  Morelos,  MEXICO,  28  Aug.  1964 
(E.  Fisher  and  D.  Verity),  in  Natural  History  Museum  of  Los 
Angeles  County.  Paratypes,  MEXICO:  13,  Carapan,  Mi- 
choacan,  1 Sept.  1962  (D.H.  Janzen;  UCB);  13,  3.4  km  NW 
Tequila,  1295  m elev.,  Jalisco,  6 Sept.  1976  (C.D.  George 
and  R.R.  Snelling;  LACM);  13,  5 mi.  W Durango,  Durango, 
21  July  1964  (J.  Powell;  UCB);  233,  25  mi.  W Hidalgo  del 
Parral,  6800  ft.  elev.,  Chihuahua,  15  July  1964  (J.A.  Chem- 
sak, J.  Powell;  UCB);  13,  “Guanajuato,  Mexique”  (Duges; 
MNHN).  UNITED  STATES:  13,  Atascosa  Mts,  4800  ft.  elev., 
5.6  mi.  W Pena  Blanca  Lake,  Santa  Cruz  Co.,  Arizona,  15 
Aug.  1974  (T.J.  Zavortink;  TJZ),  on  Acacia  angustissima, 
1145-1200. 

ETYMOLOGY 

From  Hebrew,  literally  an  enemy  or  an  evil  spirit;  in  modern 
zoological  nomenclature  an  epithet  for  a species  of  black  or 
dark  color  or  threatening  appearance. 

DISCUSSION 

The  wholly  black  pubescence  will  immediately  separate  C. 
satana  from  C.  agameta  and  C.  albiceps,  since  both  have 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  1 1 


Figures  1-8.  Male  seventh  and  eighth  sternites  and  genitalia  (ventral  and  dorsal  views),  respectively,  of:  1-4,  Centris  ( Acritocentris ) satana, 
scale  line  = 1.00  mm;  5-8,  C.  (C.)  aethiocesta.  scale  line  = 0.50  mm. 


12  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


pale  dorsal  thoracic  pubescence.  The  black,  rather  than  yel- 
low, clypeus  will  distinguish  C.  satana  from  C.  ruthannae. 
Additionally,  males  of  C.  ruthannae  have  metallic  blue  re- 
flections on  the  abdominal  terga  and  the  hairs  of  the  second 
and  third  segments  are  conspicuously  plumose. 

The  female  of  C.  satana  is  unknown,  but  very  likely  will 
be  black-haired  like  the  male  and  will  have  a similarly  black 
abdomen,  without  metallic  blue  reflections. 

Subgenus  Exallocentris  Snelling 

Centris  subg.  Exallocentris  Snelling,  1974:35.  Type-species: 
Centris  ( Melanocentris ) anomala  Snelling,  1966;  mono- 
basic and  original  designation. 

Although  I had  originally  placed  this  monotypic  subgenus 
near  Melanocentris,  Neff  and  Simpson  (1981)  have  rightly 
pointed  out  its  affinities  to  Paracentris.  In  Exallocentris  the 
elaiospathe  of  the  female  pro-  and  mesobasitarsi  is  replaced 
by  dense  pads  of  fine-branched  setae  and  the  secondary  basi- 
tibial  plate  is  sharply  marginate  and  projects  over  the  primary 
plate.  Males  differ  from  those  of  Paracentris  in  lacking 
branched  setae  on  the  gonocoxite  at  the  base  of  the  gono- 
stylus;  a conspicuous  pygidial  plate  is  present,  the  innermost 
mandibular  tooth  is  truncate,  and  the  lateral  ocellus  is  sep- 
arated from  the  inner  eye  margin  by  about  its  own  diameter. 
Males  resemble  those  of  Melanocentris,  but  the  scutellum  is 
not  bilobed  on  its  dorsal  surface  and  the  upper  inner  man- 
dibular carina  ends  near  the  base  of  the  innermost  tooth. 

Centris  ( Exallocentris ) anomala  Snelling 

Centris  ( Melanocentris ) anomala  Snelling,  1966:31-32.  6 2. 
Centris  (Exallocentris)  anomala:  Snelling,  1974:35-36  (tax.). 

This  species  has  previously  been  known  only  from  the  vi- 
cinity of  Guadalajara,  Jalisco,  Mexico.  I have  recently  seen 
a few  specimens  of  the  cleptoparasitic  bee,  Mesoplia  dugesi 
(Cockerell),  which  were  collected  with  the  type  series  of  C. 
anomala.  One  of  the  specimens  bears  the  following  note: 
“Ericrocis??/Very  evasive,  flying  low/Parasite  ??  of  the/big 
Hemisia?  not  numerous.” 

NEW  RECORDS 

MEXICO,  HIDALGO:  18,  22  mi.  SW  Actopan,  6800  ft.  elev., 
27  Aug.  1962  (Ordway  and  Marston;  UKAN).  JALISCO: 
12,  103  mi.  NE  Guadalajara,  6200  ft.  elev.,  1 Oct.  1957  (H. A. 
Scullen;  ORSU);  1033,  8 mi.  NE  Chapala,  5150  ft.  elev.,  30 
Sept.  1957  (H.A.  Scullen;  ORSU);  12,  “env.  de  Guadalajara” 
(L.  Diguet;  MNHN).  MICHOACAN:  222,  33  km  NE  Arteaga 
(Hwy  37,  km  242),  980  m elev.,  10  Nov.  1976  (E.  Fisher 
and  P.  Sullivan;  LACM).  OAXACA:  12,  9 mi.  SE  Nochixtlan, 
7 Nov.  1963  (R.F.  Smith;  UCB);  333,  Monte  Alban,  12  Oct. 
1963  (A.E.  and  M.M.  Michelbacher;  UCB).  PUEBLA:  9 33, 
“env.  de  Tehuacan”  (L.  Diguet;  MNHN).  ZACATECAS:  12, 
10  mi.  S Jalpa,  18  Sept.  1970  (R.M.  Bohart;  UCD). 

Subgenus  Xanthemisia  Moure 

Xanthemisia  Moure,  1945:401.  Type-species:  Centris  bicolor 

Lepeletier,  1841;  monobasic  and  original  designation. 


Hemisia  subg.  Xanthemisia:  Michener,  1951:2,  3,  5-6. 
Centris  subg.  Xanthemisia:  Michener,  1954:140.  Snelling, 
1974:2,  3. 

This  subgenus  was  described  for  a small  group  of  species 
characterized,  in  the  females,  by  possessing  a low,  blunt  tooth 
on  the  inner  surface  of  the  mandible,  near  the  base  of  the 
apical  tooth.  In  this  sex,  too,  the  pygidial  plate  is  abruptly 
narrowed  a short  distance  beyond  the  secondary  plate  so  that 
the  distal  portion  is  narrow  and  parallel-sided.  Males  of  Xan- 
themisia possess  giant  branched  setae  near  the  base  of  the 
gonostylus,  which  is  much  broadened  on  a vertical  plane  and 
the  maxillary  palp  is  four-segmented. 


KEY  TO  NORTH  AMERICAN  XANTHEMISIA 

la.  Male,  antenna  1 3-segmented,  basitibial  plate  absent  . . 

2 

b.  Female,  antenna  1 2-segmented,  basitibial  plate  present 

4 

2a.  Labrum  and  clypeus  wholly  black;  pubescence  of  dorsum 
of  scutellum  yellow;  erect  hairs  of  mesoscutum  longer 

than  interocellar  distance 3 

b.  Labrum,  entirely,  and  clypeus  mostly,  yellow;  pubes- 
cence of  dorsum  of  scutellum  black;  erect  hairs  of  meso- 
scutum shorter  than  interocellar  distance 

rubella  F.  Smith 

3a.  Pubescence  of  mesoscutum  wholly  yellow;  abdomen 
black  and  wholly  blackish  pubescent;  ocellocular  dis- 
tance less  than  diameter  of  anterior  ocellus  

lutea  Friese 

b.  Mesoscutum  with  interalar  band  of  blackish  pubescence; 
abdomen  reddish,  with  golden  brown  pubescence;  ocel- 
locular distance  greater  than  diameter  of  anterior  ocellus 

carolae  Snelling 

4a.  Clypeus  immaculate  and  pubescence  of  dorsum  of  scu- 
tellum yellow;  abdomen  blackish  or  dark  brown;  scopal 

hairs  dark  5 

b.  Clypeus  usually  with  a pair  of  preapical  spots;  pubes- 
cence of  dorsum  of  scutellum  dark  brown;  abdomen  dull 

ferruginous;  scopal  hairs  pale  rubella  F.  Smith 

5a.  Mesoscutum  wholly  yellow  pubescent;  abdomen  black- 
ish, with  blackish  pubescence;  first  flagellar  segment 
longer  than  following  three  combined  ....  lutea  Friese 
b.  Mesoscutum  wholly  blackish  pubescent;  abdomen  dark 
brown  with  reddened  apical  zone  and  yellowish  to  yel- 
lowish red  pubescence;  first  flagellar  segment  no  longer 
than  following  three  combined  carolae  Snelling 

Centris  ( Xanthemisia ) carolae  Snelling 

Centris  ( Xanthemisia ) carolae  Snelling,  1966:24-25.  3. 

This  species  has  been  previously  known  only  from  the  unique 
male  type  from  Tuxtla  Chico,  Chiapas,  Mexico.  A few  fe- 
males are  now  available.  They  may  be  separated  from  our 
other  species  by  the  largely  blackish  pubescence  on  the  tho- 
racic dorsum,  yellowish  only  on  the  scutellum  and  meta- 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  13 


notum.  The  pubescence  of  the  abdomen  is  dull  reddish  over 
the  brownish  integument. 

NEW  RECORDS 

EL  SALVADOR:  29$,  Cerro  Verde,  6800  ft.  elev.,  29  June 
1963  (M.E.  Irwin  and  D.Q.  Cavagnaro;  UCB);  1$,  same  data 
(CAS).  COSTA  RICA:  1$,  “El  Fuente,”  8 July  1937  (A.  Al- 
faro No.  177;  AMNH). 

Centris  ( Xanthemisia ) lutea  Friese 

Centris  lutea  Friese,  1899:43.  <3  9. 

Centris  (Xanthemisia)  lutea : Michener,  1954:140  (distr.). 

Among  the  North  American  Centris,  this  species  is  easily 
known  by  the  wholly  blackish  pubescence,  except  that  of  the 
thoracic  dorsum  which  is  bright  lemon-yellow.  Michener 
(1954)  recorded  this  primarily  South  American  species  from 
Panama. 

NEW  RECORDS 

MEXICO,  CHIAPAS:  2<3<3,  46  km  N Chilpacingo,  580  m 
elev.,  4-6  Aug.  1977  (E.M.  Fisher  and  P.  Sullivan;  LACM). 
JALISCO:  1<3,  48  mi.  N Guadalajara,  8 Sept.  1966  (R.J. 
Hamton;  LACM).  OAXACA:  1<3,  20  mi.  E El  Camaron,  21 
July  1956  (J.W.  MacSwain;  UCB).  COSTA  RICA,  GUA- 
NACASTE:  19,  [Hacienda]  Comelco,  8 km  NW  Bagaces,  21 
Jan.  1 972  (P.A.  Opler;  UCB),  on  Bvrsonima  sp.;  19,  Hacienda 
Comelco,  24  km  NW  Canas,  7 Feb.  1972  (E.R.  Heithaus; 
LACM),  on  Bvrsonima  crassifolia,  0730-0830.  SAN  JOSE: 
19,  San  Jose,  no  further  data  (USNM). 

Centris  ( Xanthemisia ) rubella  F.  Smith 

Centris  rubella  F.  Smith,  1854:372.  9. 

Hemisia  ( Xanthemisia ) rubella:  Michener,  1951:6  (tax.). 

The  ferruginous  abdomen  and  maculate  female  clypeus  will 
readily  separate  C.  rubella  from  all  other  species  of  Xanthe- 
mesia.  This  species  has  previously  been  known  only  from 
South  America. 

NEW  RECORDS 

PANAMA,  CANAL  ZONE:  19,  Barro  Colorado  Island,  19 
Aug.  1968  (L.S.  Kimsey;  LACM),  “wood  nester”;  399,  same 
locality,  27  Apr.,  3 May,  5 May  1980  (K.E.  Steiner;  LACM, 
UCD),  on  Byrsonima  crassifolia ; 19,  Gatun,  20  May  1980 
(K.E.  Steiner;  UCD),  on  B.  crassifolia ; 399,  same  locality,  30 
Oct.  and  3 Nov.  1977  (K.E.  Steiner;  UCD),  on  Stigmaphyllon 
hypargyreum. 

Subgenus  Centris  Fabricius 

Centris  Fabricius,  1804:354.  Type-species:  Apis  haemor- 
rhoidalis  Fabricius,  1775;  designated  by  Internatl.  Comm. 
Zool.  Nomencl.,  op.  567,  1959. 


Hemisia  Klug,  1807:227.  Type-species:  Apis  haemorrhoi- 
da/is  Fabricius,  1 775;  designated  by  Cockerell,  1906. 
Centris  subg.  Cyanocentris  Friese,  1900b:243.  Type-species: 
Apis  versicolor  Fabricius,  1775;  designated  by  Sandhouse, 
1943. 

Centris  subg.  Poecilocentris  Friese,  1900b:244.  Type-species: 
(Centris  [Poecilocentris]  fasciateIla  Friese,  1900)  = Centris 
fasciata  F.  Smith,  1854;  designated  by  Sandhouse,  1943. 

Species  in  this  subgenus  normally  have  the  abdomen  metallic 
blue  or  blue-green  in  both  sexes  and  with  pale  tergal  maculae 
in  the  male  and  often  in  the  female  as  well.  In  a few  species, 
such  as  C.  inermis  Friese,  the  abdomen  is  largely  ferruginous 
and  in  others,  such  as  C.  eisenii  W.  Fox,  both  sexes  have 
much  of  the  terga  taken  up  by  yellow  bands.  In  both  sexes, 
too,  the  mandibles,  labrum,  and  clypeus  are  conspicuously 
maculate,  the  clypeal  maculation  of  the  female  usually  in  an 
inverted  T-shape  or,  more  rarely,  an  inverted  Y-shape.  Both 
sexes  have  five-segmented  maxillary  palps,  the  female  with 
slender,  usually  quadridentate,  mandible,  and  a distinct  sec- 
ondary basitibial  plate  with  an  overhanging  margin.  Males 
possess  giant  branched  setae  along  the  inner  margin  of  the 
styliform  apical  process  of  gonocoxite. 

Some  of  the  most  exasperating  taxonomic  problems  in 
Centris  are  encountered  in  the  nominate  subgenus.  Many  of 
the  species  are  widely  distributed  and  may  be  locally  abun- 
dant. The  more  widely  distributed  species  tend  to  exhibit  a 
variety  of  phenotypes  throughout  their  ranges,  phenotypes 
that  are  often  strikingly  different  from  one  another.  Many  of 
these  variant  populations  have  been  given  formal  names.  As 
a rule,  these  divergent  populations  represent  selected  points 
along  a dine  of  variation  and  it  is  impossible  to  fix  to  any 
of  these  a discrete  distribution  and  set  of  identifying  char- 
acteristics. 

The  situation  becomes  more  complex  when  a species  is 
distributed  through  the  islands  of  the  Caribbean.  Since  the 
ranges  are  disjunct,  the  various  insular  populations  tend  to 
breed  true.  Even  here,  however,  there  usually  are  discrete 
clinal  trends,  proceeding  from  one  end  of  the  island  distri- 
bution to  the  other.  But,  a great  deal  of  collecting  must  be 
done  in  the  Caribees  before  any  understanding  can  be  gained 
of  these  forms. 

Further  complicating  the  taxonomy  of  this  subgenus  is  the 
fact  that  some  species  perhaps  most,  possess  metanders,  i.e., 
males  which  are  unusually  large  and  robust,  with  very  stout 
legs,  and  much  more  extensively  maculate  than  normal  males. 
Metanders  may  be  easily  assumed  to  represent  a different 
species.  In  general,  all  large,  robust,  richly  marked  males 
should  be  assumed  to  be  probable  metanders. 

Yet  another  difficulty  is  that  at  least  one  species  (C.  iner- 
mis) is  dichromatic  in  both  sexes.  There  is  a color  phase  with 
red  abdomen  and  one  with  metallic  blue-green  abdomen. 
There  is  some  evidence  that  one  phase  tends  to  fly  earlier 
than  the  other,  but  there  is  a broad  temporal  overlap.  Not 
surprisingly,  the  two  color  phases  have  been  thought  to  be 
separate  species.  There  are  very  likely  more  such  cases  in 
this  subgenus. 

The  following  key  is  for  those  species  which  occur  in  Cen- 


14  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


tral  America;  it  does  not  include  C.  errans  W.  Fox,  a species 
found  only  in  Florida. 

KEY  TO  CENTRAL  AMERICAN  CENTRIS,  S.  S I R. 

1 a.  Male,  antenna  1 3-segmented  and  basitibial  plate  absent 

2 

b.  Female,  antenna  12-segmented  and  basitibial  plate 

present  9 

2a.  Disc  of  fourth  tergum  with  simple,  usually  blackish, 
hairs  only  (plumose  hairs  may  be  present  on  apical 

zone)  3 

b.  Disc  of  fourth  tergum  with  abundant,  usually  whitish, 
suberect  plumose  hairs  which  extend  onto  apical  zone 
(may  be  only  a band  across  segment  basad  of  apical 

zone,  often  extended  cephalad  in  middle)  7 

3a.  Second  to  sixth  abdominal  terga  with  broad  preapical 
yellow  bands  which  cover  most  of  each  segment,  nar- 
row basal  area  bluish  to  rufescent,  apical  margin  trans- 
parent; sterna  largely  rufescent  4 

b.  Second  to  sixth  abdominal  terga  largely  bluish  or  fer- 
ruginous, second  usually  with  lateral  spots  only,  fol- 
lowing segments  with  lateral  spots,  narrow  transverse 
bands  (often  interrupted  and  usually  hidden  under  mar- 
gin of  preceding  segment)  or  immaculate;  if  bands  are 
present,  they  are  dull  reddish,  usually  interrupted  in 
middle,  and  fourth  tergum  with  broad  area  along  distal 

margin  with  abundant  plumose  hairs  5 

4a.  Facial  marks  whitish;  first  flagellar  segment  at  least 
1.25  x scape  length;  hairs  along  apical  band  of  fourth 

tergum  dark,  mostly  simple  eisenii  W.  Fox 

b.  Facial  marks  distinctly  yellowish;  first  flagellar  segment 
about  as  long  as  scape,  rarely  as  much  as  1.1  x scape 
length;  apical  band  of  fourth  tergum  with  abundant 

plumose  pale  hairs  aethyctera  Snelling 

5a.  Ground  color  of  abdominal  segments  ferruginous,  sec- 
ond segment  maculate  at  side,  rarely  with  a complete 
or  narrowly  interrupted  subbasal  band;  transverse  bands 
of  remaining  segments,  if  present,  hidden  under  margin 

of  preceding  segment  6 

b.  Ground  color  of  first  terga,  and  of  sterna,  bluish,  sub- 
lateral spots  or  narrowly  interrupted  bands  on  one  or 
more  segments;  rarely,  second  to  sixth  segments  with 
broad,  transverse,  median  yellowish  or  rufescent  bands 

inermis  Friese  (part) 

6a.  Flairs  of  thoracic  dorsum  black-tipped  

obscurior  Michener 

b.  Hairs  of  thoracic  dorsum  uniformly  ochreous  

inermis  Friese  (part) 

7a.  Pubescence  of  thoracic  dorsum  uniformly  ochreous  to 
ferruginous,  no  blackish  hairs  present;  smaller  species, 
head  width  less  than  6. 1 mm  and  usually  less  than  6.0 

mm;  (metander  unknown) 8 

b.  Pubescence  of  thoracic  dorsum  pale  anteriorly  and  pos- 
teriorly, a broad  interalar  brown  to  black  band  present, 
often  replacing  much  pale  pubescence,  especially  on 
scutellum;  larger  species,  head  width  at  least  5.5  mm 
and  usually  over  6.3  mm  (metander  larger,  robust,  legs 


and  abdomen  abundantly  yellow  maculate  or  both  areas 
largely  ferruginous  with  limited  yellow  maculae)  . . 

flavifrons  group  (see  Text) 

8a.  Apical  two  or  three  abdominal  terga  reddish;  mandible 
mostly  yellowish  on  outer  face;  pubescence  of  thoracic 

dorsum  usually  ochreous  adani  Cockerell 

b.  Fourth  and  fifth,  usually  sixth,  abdominal  tergites  blue, 
except  colorless  margins;  mandible  with  small  pale  spot 
near  base;  pubescence  of  thoracic  dorsum  usually  dark 

ferruginous  decolorata  Lepeletier 

9a.  Terga  2-5  each  with  entire  transverse  yellow  fascia 

10 

b.  Terga  2-5  without  yellow  fasciae  11 

10a.  Hairs  of  thoracic  dorsum  black-tipped;  scape  yellow 
beneath;  fourth  tergum  with  erect,  plumose,  white  hairs; 

HW  less  than  6.0  mm  aethyctera  Snelling 

b.  Hairs  of  thoracic  dorsum  not  black-tipped;  scape  im- 
maculate beneath;  fourth  tergum  with  simple  hairs  only; 

HW  at  least  6.3  mm  eisenii  Fox 

1 la.  Thoracic  dorsum  with  conspicuous  interalar  band  of 
black  pubescence  which  may  cover  much  of  mesoscu- 
tum  (sometimes  reduced  to  median  patch),  sharply 

contrasting  with  pale  scutellar  hairs  12 

b.  Thoracic  dorsum  without  interalar  band  of  black  pu- 
bescence; scutellar  hairs  not  contrastingly  colored 

14 

12a.  Larger  species,  head  width  greater  than  6.5  mm,  but  if 
as  little  as  6.25  mm,  scopa  is  principally  black;  disc  of 
clypeus,  in  profile,  distinctly  convex  basad  and  flat- 
tened distad  (Fig.  16)  13 

b.  Smaller  species,  head  width  5.95-6.25  mm;  scopa  pale, 
with  golden  brown  hairs  apicad  on  metabitarsus;  disc 
of  clypeus,  in  profile,  very  weakly  convex  basad  (Fig. 

15)  aethiocesta,  new  species 

13a.  Larger  species,  head  width  7.17-7.7  mm;  scopa  with 
pale  hairs  at  least  anterobasally  on  metatibia,  often 
largely  pale;  pygidial  plate  and  basitibial  plate  as  in 

Figs.  38  and  29  flavifrons  (Fabricius) 

b.  Smaller  species,  head  width  6.27-7.2  mm;  scopa  black, 
a few  posteroapical  hairs  on  metabasitarsus  pale  at  tips; 
pygidial  and  basitibial  plates  as  in  Figs.  39  and  30 

flavofasciata  Friese 

14a.  Abdomen  mostly  ferruginous,  one  or  more  terga  often 
partially  bluish  or  greenish  toward  apical  margin  ..15 
b.  At  least  first  three  terga  wholly  dark  blue  or  blue-green 

17 

15a.  Hairs  of  thoracic  dorsum  not  dark-tipped;  if  slightly 
tipped,  terga  2-4  with  conspicuous  apical  hair  bands; 

erect  plumose  hairs  of  tergum  4 pale  16 

b.  Hairs  of  thoracic  dorsum  conspicuously  dark-tipped 
and  terga  2-4  without  apical  hair  bands;  erect  plumose 

hairs  of  tergum  4 dark  obscurior  Michener 

16a.  Terga  2-4  with  conspicuous  apical  hair  bands;  basal 
margin  of  labrum  convex  across  entire  width,  convex 
ridge  with  fine,  widely  spaced  longitudinal  ridges;  ex- 
ternal stripe  of  protibia  not  reaching  apical  three-fourths 

of  segment meaculpa,  new  name 

b.  Terga  2-4  without  apical  hair  bands;  basal  margin  of 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  15 


labrum  smooth  and  flat  between  punctures;  external 
stripe  of  protibia  reaching  three-fourths,  or  more,  of 

tibial  length inermis  Friese  (part) 

17a.  Apical  margin  of  tergum  4 and  all  of  tergum  5 ferru- 
ginous; scopa  entirely  pale  or  entirely  dark;  terga  2-3 

with  or  without  apical  hair  bands  18 

b.  Terga  4 and  5 blue-green;  scopa  pale,  with  brownish 
setae  posteriorly  and  apically  on  basitarsus;  terga  2-3 

without  apical  hair  bands  decolorata  Lepeletier 

18a.  Scopa  black;  terga  2-3  without  apical  hair  bands;  ex- 
ternal stripe  of  protibia  extending  half,  or  more,  length 

of  segment inermis  Friese  (part) 

b.  Scopa  pale;  terga  2-3  with  apical  hair  bands,  that  of  2 
interrupted  in  middle;  external  stripe  of  protibia  less 

than  half  length  of  segment,  often  absent 

adani  Cockerell 


Centris  ( Centris ) aethyctera  Snelling 

Centris  ( Centris ) aethyctera  Snelling,  1974:23-26.  $ 2. 

Centris  aethyctera  is  a common  species  ranging  from  Mexico 
to  Panama.  It  can  only  be  confused  with  C.  eisenii  in  this 
region,  a larger  species  with  the  hairs  of  the  thoracic  dorsum 
without  black  tips.  In  previous  literature  it  has  been  confused 
with  the  Antillean  species,  C.fasciata  F.  Smith  (see  Snelling, 
1974),  and  all  records  of  C.  fasciata  (or  its  synonym  C.fas- 
ciatella  Friese)  in  Central  America  seem  to  be  based  on  C. 
aethyctera. 

In  Panamanian  populations  of  C.  aethyctera  the  bands  of 
the  abdominal  terga  are  conspicuously  shorter.  The  abdomen 
thus  appears  to  be  principally  ferruginous,  with  short,  trans- 
verse yellow  fasciae  across  the  disc  of  each  segment. 

Centris  ( Centris ) decolorata  Lepeletier 

Centris  decolorata  Lepeletier,  1 84 1 : 1 60.  <3. 

Centris  ( Cyanocentris ) decolorata:  Friese,  1900b:243,  325.  3 
(not  2?). 

Centris  versicolor:  Cheesman,  1929:142.  Misidentification. 
Centris  (Centris)  obscuriventris:  Michener,  1954:138.  Mis- 
identification. 

Centris  ( Centris ) decolorata:  Snelling,  1966:23  (distr. ). 

Friese  (1900b)  associated  females  with  C.  decolorata.  but 
noted  that  the  association  might  be  incorrect.  In  the  event 
that  this  should  prove  to  be  true,  he  proposed  that  the  females 
be  called  C.  obscuriventris.  Whether  or  not  his  female  spec- 
imens actually  are  those  of  C.  decolorata  can  only  be  deter- 
mined after  his  specimens  are  examined.  In  the  meantime, 
C.  obscuriventris  must  be  considered  a name  of  dubious  va- 
lidity and  identity. 

There  is  no  doubt,  however,  that  Central  American  spec- 
imens, previously  identified  as  C.  obscuriventris.  are  conspe- 
cific  with  C.  decolorata.  a common  species  throughout  the 
Caribbean  islands,  coastal  northern  South  America,  and  east- 
ern Central  America.  The  range  extends  north  to  southern 


Texas,  but  C.  decolorata  appears  to  be  absent  from  Florida, 
even  though  present  on  Cuba.  The  record  from  Cristobal, 
Canal  Zone,  Panama,  of  C.  versicolor  by  Cheesman  (1929) 
is  based  on  misidentified  specimens  of  C.  decolorata. 


Centris  ( Centris ) errans  W.  Fox 

Centris  errans  W.  Fox,  1899:65.  2. 

Centris  versicolor:  Lutz  and  Cockerell,  1920:561  (in  part); 

Mitchell,  1962:335-336  (misidentification). 

Centris  ( Centris ) versicolor:  Hurd,  1979:2175  (in  part,  mis- 
identification). 

This  species  occurs  in  Florida  and  has  often  been  cited  as  C. 
versicolor  (Fabricius,  1775).  However,  Moure  (1960b)  ex- 
amined the  type  of  C.  versicolor  and  redescribed  the  type 
specimen;  the  true  C.  versicolor  is  “probably  confined  to 
lesser  Antilles,”  according  to  Moure.  Until  all  the  represen- 
tatives of  this  very  difficult  complex  can  be  reexamined,  it 
seems  best  to  follow  Moure’s  restricted  interpretation  of  C. 
versicolor  and  to  regard  C.  errans  as  a distinct  species. 

Centris  ( Centris ) inermis  Friese 

Centris  inermis  Friese,  1899:46.  3 2. 

Centris  poecila  var.  segregata  Crawford,  1906:159.  2.  NEW 
SYNONYMY. 

Centris  inermis  var.  gualanensis  Cockerell,  1912:568.  3.  2. 
Centris  (Rhodocentris)  robusta  Cockerell,  1949:478-479.  3. 
NEW  SYNONYMY. 

Centris  inermis  subsp.  pallidifrons  Cockerell,  1949:479.  3. 
Centris  ( Centris ) inermis:  Snelling,  1974:30  (syn.,  var.). 
Centris  ( Centris ) segregata:  Snelling,  1974:34-35  (tax.,  distr.). 

This  is  a common  Central  American  species,  ranging  from 
Mexico  to  Panama  and  into  northern  South  America.  It  is 
also  a very  unusual  species,  one  which  suggests  that  the  tax- 
onomy of  the  nominate  subgenus  may  be  fraught  with  more 
complexities  than  was  previously  supposed.  Males  of  C.  iner- 
mis are  morphologically  bimodal;  there  is  a metandric  form, 
very  different  in  appearance  from  the  normal  males.  Addi- 
tionally, however,  there  are  two  very  different  color  pheno- 
types in  both  sexes. 

The  nominate  phenotype,  in  both  sexes,  has  the  legs  and 
abdomen  ferruginous;  in  females  there  is  usually  a conspic- 
uous patch  of  metallic  blue  across  the  base  of  the  fourth 
tergite.  The  second  phenotype  was  described  as  C.  poecila 
var.  segregata  and  later  considered  to  be  a separate  species. 
In  this  form  the  legs  are  blackish  brown  and  the  first  four 
abdominal  segments  are  dark  metallic  blue. 

The  recognition  of  C.  segregata  as  a synonym  of  C.  inermis 
is  possible  thanks  to  the  observations  of  R.  Coville  and  G. 
Frankie.  They  have  found  that  emergent  bees  from  nests 
provisioned  by  C.  inermis  females  were,  sometimes,  the  blue- 
abdomen  form,  C.  segregata.  This  fact,  coupled  with  the  lack 
of  morphological  differences  between  the  two  forms,  is  the 
basis  for  the  current  synonymy.  Additional  support  lies  in 


16  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


the  complete  allopatry  of  the  two  forms.  Frankie  and  Coville 
will  publish  their  observations  separately. 

Centris  {Cent r is)  meaculpa,  new  name 

Figure  33 

Centris  ( Cyanocentris ) nitens:  Friese,  1900b:330.  2 (in  part; 
misidentification). 

Centris  (Centris)  emhescens  Snelling,  1974:27-28.  2.  Preoc- 
cupied. 

My  C.  erubescens  is  a junior  homonym  of  C.  costaricensis 
var.  erubescens  Friese,  1925,  a synonym  of  C.  vittata  Le- 
peletier.  The  new  specific  epithet  is,  of  course,  the  Latin 
phrase  for  “my  fault”  and  seems  apropos  in  this  case. 

In  some  of  the  specimens  recorded  below,  the  abdominal 
tergites,  especially  the  second  to  fourth  segments,  are  exten- 
sively bluish,  rather  than  wholly  ferruginous.  They  thus 
somewhat  resemble  females  of  C.  adani.  However,  in  C. 
adani  the  erect  hairs  of  the  the  fourth  tergite  are  longer  and 
are  plumose  at  their  tips,  as  noted  in  the  key.  In  most  spec- 
imens of  C.  meaculpa  the  clypeus  is  more  or  less  distinctly 
obliquely  rugulose,  the  rugulae  directed  toward  the  apical 
middle  portion  of  the  segment.  The  clypeus  of  C.  adani  tends 
to  be  smooth  between  well  spaced  punctures. 

The  female  which  Friese  (1900b)  recorded  from  Mexico 
as  C.  nitens  Lepeletier,  a Brazilian  species,  is  misidentified. 
The  specimen  is  in  the  Paris  Museum  and  has  been  made 
available  to  me;  it  is  a female  of  C.  meaculpa  with  the  first 
three  abdominal  terga  primarily  metallic  bluish  and  the  spec- 
imen agrees  well  with  the  characteristics  of  this  species,  es- 
pecially in  the  structure  of  the  labrum  and  of  the  basitibial 
plate. 

When  I originally  described  this  species,  I was  not  fully 
aware  of  its  apparent  affinities  with  a small  group  of  primarily 
Brazilian  species.  Included  in  this  group  are  C.  aenea  Le- 
peletier, 1841,  C.  caixensis  Ducke,  1907,  and  C.  nitens  Le- 
peletier, 1841.  Females  of  this  group  are  similiar  in  that  the 
pubescence  of  the  thorax  is  ochreous  (paler  on  the  sides),  the 
abdominal  terga,  except  the  fifth  and  sixth,  are  metallic  green- 
ish to  bluish,  the  discs  of  the  second  and  third  segments 
possess  abundant  appressed  simple  hairs,  and  the  margins 
of  the  second  and  third  segments  have  a definite  fascia  of 
appressed  pale  hairs  which  may  be  plumose.  In  these  Bra- 
zilian species  the  basal  margin  of  the  labrum  is  flat  and  is 
smooth  and  shiny  between  sparse  punctures.  The  labrum  of 
C.  meaculpa  has  a definite  convexity  across  the  basal  margin 
and  the  convex  area  is  crossed  by  widely  spaced  longitudinal 
ridges. 

Of  the  Brazilian  species,  C.  caixensis  appears  to  be  the 
most  distinct.  I have  seen  a single  female,  marked  as  “Type,” 
from  the  collections  of  the  Paris  Museum:  Maranhao,  Caixas, 
30  June  1907,  collected  by  A.  Ducke.  The  second  to  fifth 
terga  each  bear  a small,  lateral,  pale  yellow  mark;  the  basi- 
tibial plate  (Fig.  32)  is  short,  and  the  secondary  plate  is  short, 
broad,  and  convexly  transverse.  In  all  of  the  other  species, 
including  C.  meaculpa  (Fig.  33),  the  secondary  plate  is  elon- 
gate, with  its  anterior  margin  approximately  parallel  with. 


and  well  removed  from,  the  anterior  margin  of  the  primary 
plate.  The  fascial  hairs  of  the  second  and  third  terga  of  C. 
caixensis  are  yellow  and  simple. 

Of  the  two  remaining  species,  C.  aenea  may  be  recognized 
by  the  golden  or  somewhat  coppery  color  of  the  discal  and 
fascial  hairs  of  the  second  and  third  terga.  In  C.  nitens  the 
discal  hairs  are  fuscous,  sharply  contrasting  with  the  golden 
to  coppery  color  of  the  fascial  hairs.  The  secondary  pygidial 
plate  is  more  abruptly  narrowed  in  C.  nitens , which  is  the 
smaller  of  the  two  species. 

This  species  was  described  from  two  females  67  km  E 
Escarcega,  Campeche,  Mexico.  A few  additional  specimens 
are  now  available. 

NEW  RECORDS 

MEXICO,  TAMA  ULIPAS:  12,  8 mi.  NW  Nuevo  Morelos, 
22  July  1962  (Univ.  Kans.  Mex.  Exped.;  UKAN),  on  Kall- 
stroemia  sp.  SAN  LUIS  POTOSP.  12,  El  Salto,  1800  ft.  elev., 
8 June  1961  (Univ.  Kans.  Mex.  Exped.;  UKAN);  12,  El  Salto 
(above  falls),  21  July  1962  (Ordway  and  Roberts;  UKAN), 
on  Leguminoseae.  VENA  CRUZ:  12,  22  mi.  SE  Jalapa,  1 100 
ft.  elev.,  29  June  1953  (Univ.  Kans.  Mex.  Exped.;  UKAN); 
222,  3 mi.  SW  Paso  del  Toro,  50  ft.  elev.,  23  June  1961 
(Univ.  Kans.  Mex.  Exped.;  UKAN).  YUCATAN:  12,  Piste, 
12  June  1967  (E.C.  Welling;  LACM).  STATE  UNKNOWN: 
12,  “Mexique,  Cote  Occid.,”  no  date  except  1864  (L.  Biart; 
MNHN). 

Centris  ( Centris ) obscurior  Michener 
Centris  (Centris)  obscurior  Michener,  1954:138-140.  2 6. 
This  species  has  previously  been  reported  only  from  Panama. 

NEW  RECORDS 

COSTA  RICA,  HEREDIA:  12,  Puerto  Viejo,  Sarapiqui,  30 
July  1965  (D.H.  Janzen;  UKAN);  822,  37<$<5,  Finca  La  Selva, 
near  Puerto  Viejo,  6 May  to  27  July  (D.R.  Perry;  LACM), 
on  Dipteryx  panamensis,  Dussia  sp.,  Hymenolobium  sp., 
Byrsonima  sp.,  and  Vochysia  sp.  PUNTARENAS:  12,  6 km 
S San  Vito,  19-21  Mar.  1967  (UKAN).  SAN  JOSE:  222,  San 
Jose,  1160  m elev.,  17  July  1964  (M.  Naumann;  UKAN), 
on  Solatium  sp.;  222,  San  Jose,  1160  m elev.,  8 June  1963 
(C.D.  and  D.R.  Michener;  UKAN),  on  Solatium  wendlandy 
222,  Ciudad  Universitaria,  San  Jose,  28  July  1965  (S.J.  Ar- 
nold; UCB),  on  Duranta  repens\  1622,  1 mi.  ESE  San  Isidro 
de  General,  21  July  1965  (S.J.  Arnold;  UCB),  on  Rhyn- 
ch  a nth  era  mexicana,  0815-0950.  MEXICO,  CHIAPAS:  12, 
Simojovel,  1-16  Aug.  1958  (J.A.  Chemsak;  UCB);  12,  Santo 
Domingo,  15  mi.  S Simojovel,  8 July  1958  (J.A.  Chemsak; 
UCB);  3<5<5,  3 mi.  SE  San  Juan  del  Bosque,  16  Aug.  1958 
(J.A.  Chemsak;  UCB);  12,  Yaxoquintela,  560  m elev.,  30 
Aug.  1978  (J.E.  Rawlons;  CORN). 

Centris  (Centris)  flavifrons  Group 

To  this  group  are  assigned  a number  of  medium-sized  to 
large  species  in  which  the  integument  of  the  head,  thorax. 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  17 


and  legs  is  blackish  with  limited  whitish  or  yellowish  marks 
on  the  face  and  legs.  The  abdomen  is  mainly  metallic  blue 
to  blue  green,  in  the  males  with  yellowish  maculations,  at 
least  in  the  second  tergite;  the  apical  segments  may  be  red- 
dish. In  both  sexes  the  thorax  is  either  largely  cinereous  pu- 
bescent and  with  a broad  interalar  band  of  blackish  hairs,  or 
principally  dark  pubescent  but  with  the  scutellum  and  meta- 
notum  pale  pubescent.  The  wings  are  light  brownish  to  black- 
ish. 

Morphologically,  the  species  tend  to  be  very  similar  and 
species  limits  are,  at  present,  very  poorly  understood.  The 
following  key  will  serve  to  separate  those  species  which  I am 
presently  able  to  recognize. 

KEY  TO  SPECIES,  C.  FLA  VIFRONS  GROUP 

la.  Male,  antenna  13-segmented  and  basitibial  plate  absent 

2 

b.  Female,  antenna  1 2-segmented  and  basitibial  plate  pres- 
ent   6 

2a.  Ocellocular  distance  no  more  than  0.90  times  diameter 
of  anterior  ocellus  and  usually  less;  lower  inner  man- 
dibular carina  ending  in  a small,  obscure  tooth-like  pro- 
cess (Fig.  13);  smaller  species,  head  width  less  than  5.8 

mm 3 

b.  Ocellocular  distance  at  least  0.95  times  diameter  of  an- 
terior ocellus  and  often  greater;  lower  inner  mandibular 
carina  often  terminating  in  a prominent  tooth-like  pro- 
cess (Fig.  14);  usually  larger  species,  head  width  usually 
more  than  6.0  mm,  but  may  be  a little  as  4.8  mm  . . 4 
3a.  Clypeus  moderately  shiny,  punctures  fine  and  close,  in- 
terspaces very  finely  roughened;  apical  margin  of  third 
tergite  with  transverse  band  of  pale  yellowish  hairs;  pu- 
bescence of  side  of  propodeum  pale  (Ecuador,  Peru) 

buchwaldi  Friese 

b.  Clypeus  shiny  and  subpolished  between  fine,  close  punc- 
tures, except  in  broad,  impunctate  median  line;  third 
tergum  blackish  pubescent  along  apical  margin;  side  of 
propodeum  brown  pubescent  (El  Salvador,  Costa  Rica, 

Panama)  aethiocesta,  new  species 

4a.  Larger  species,  head  width  always  over  5.2  mm  and 
usually  greater  than  6.0  mm;  clypeal  disc,  at  least  in  large 
part,  shiny  and  weakly,  or  not  at  all,  tessellate;  second 
tergum,  at  least,  with  transverse  yellow  fascia  across  base, 
sometimes  narrowly  interrupted  in  middle  (widespread) 

5 

b.  Smaller  species,  head  width  less  than  5.0  mm;  clypeal 
disc  moderately  shiny,  conspicuously  tessellate  and 
roughened,  more  weakly  so  along  midline;  second  ter- 
gum with  a pair  of  widely  separated  spots  (Eucador) 

nigrofasciata  Friese 

5a.  Lower  inner  mandibular  carina  ending  in  a long,  acute 
tooth-like  process.  Normal  male  with  broad,  basal,  yel- 
low fasciae  across  base  of  second  to  fourth  terga  which 
are  complete  or  narrowly  interrupted,  that  of  second 
nearly  as  long  as  apical  area  beyond  it;  punctures  of 
clypeal  disc  numerous  and  close,  interspaces  more  or 
less  distinctly  tessellate,  especially  basad.  Metander  with 


clypeus  slightly  roughened,  punctures  close  to  sparse; 
clypeal  disc  entirely  yellow  (S  Arizona  to  Panama)  . . . 

flavofasciata  Friese 

b.  Lower  inner  mandibular  carina  ending  in  low,  obtusely 
triangular  process  (Fig.  14).  Normal  male  with  narrow 
basal  fasciae  on  second  to  fourth  terga,  that  of  second 
often  broadly  interrupted  in  middle,  or,  when  complete, 
distinctly  shorter  in  middle  than  apical  area  beyond  it; 
clypeal  disc  smooth  and  polished,  punctures  at  side  sparse. 
Metander  with  clypeus  smooth  and  polished,  punctures 
sparse  to  scattered;  laterobasal  black  mark  of  clypeus 
extending  onto  disc  (widespread) . .flavifrons  (Fabricius) 
6a.  Apical  margin  of  third  tergite  with  a narrow  band  of 
white  or  yellowish  red  hairs  which  sharply  contrast  with 
the  black  discal  hairs,  the  pale  hairs  closer  than  those  of 

disc  and  projecting  well  beyond  margin  7 

b.  Apical  margin  of  third  tergite,  across  middle  one-half 
with  hairs  short,  black,  no  closer  than  short,  black  hairs 
of  disc  and  hardly  projecting  beyond  margin  of  segment 

' 8 

7a.  Scopa  pale;  marginal  hairs  of  third  tergite  whitish,  straight; 
scape  pale  beneath;  fifth  tergite  wholly  ferruginous  (Ec- 
uador)   nigrofasciata  Friese 

b.  Scopa  dark;  marginal  hairs  of  third  tergite  yellowish  red, 
their  tips  curved  laterad;  scape  dark;  fifth  tergite  mostly 
metallic  blue-green  (Ecuador,  Peru)  . . buchwaldi  Friese 
8a.  Head  width  usually  exceeding  6.7  mm,  rarely  as  little  as 
6.2  mm;  scopa  usually  mostly  dark;  clypeal  punctures 
moderate  in  size  and  on  side  of  disc  separated  by  a 
puncture  diameter  or  more;  hairs  of  side  of  propodeum 

pale,  but  if  dark,  pleural  hairs  also  dark 9 

b.  Head  width  less  than  6.3  mm;  scopa  pale;  clypeal  punc- 
tures fine  and  close;  pleura  with  pale  hairs  and  side  of 
propodeum  with  brown  hairs  (El  Salvador,  Costa  Rica, 

Panama)  aethiocesta,  new  species 

9a.  Larger  species,  head  width  7.0-7. 7 mm;  scopa  with  pale 
hairs,  at  least  anterobasally  on  metatibia,  often  largely 
pale;  pygidial  and  basitibial  plates  as  in  Figs.  38  and  29 

(widespread)  flavifrons  (Fabricius) 

b.  Smaller  species,  head  width  6. 2-7. 2 mm;  scopa  black,  a 
few  pale-tipped  hairs  posteroapically  on  metabasitarsus; 
pygidial  and  basitibial  plates  as  in  Figs.  39  and  30  (S 
Arizona  to  Panama)  flavofasciata  Friese 

Centris  ( Centris ) aethiocesta,  new  species 

Figures  5-8,  13,  15,  31,  40 

Centris  ( Centris ) nigrofasciata:  Michener,  1954:138.  Mis- 
identification. 

DIAGNOSIS 

Medium-sized  species  with  thoracic  pubescence  whitish,  but 
with  black  interalar  band  and  brown  pubescence  on  propo- 
deal  side;  abdomen  metallic  blue-green.  Female  with  pale 
scopa,  broad  basitibial  plate,  and  clypeal  disc  flattened  in 
profile.  Male  with  clypeal  disc  flattened  in  profile;  abdominal 
terga  4-7  metallic  green,  immaculate;  punctures  of  clypeal 


18  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


disc  numerous  and  mostly  separated  by  about  a puncture 
diameter. 

DESCRIPTION 

HOLOTYPE  FEMALE.  Measurements  (mm).  Head  width 
5.54  (5.95-6.26);  head  length  4.26  (3.95-4.21);  wing  length 
1 1.0  (1 1.0-12.0);  total  length  15.5  (16.6-18.0). 

Head.  1 .43  ( 1 .49-1 .53)  times  broader  than  long;  occipital 
margin  nearly  flat  in  frontal  view  and  slightly  below  level  of 
tops  of  eyes;  ocelli  anterior  to  occipital  margin;  inner  orbits 
moderately  convergent  above,  upper  frontal  width  0.90  (0.89- 
0.92)  times  lower  frontal  width.  Mandible  slender,  tridentate, 
apical  margin  oblique.  Labrum  about  twice  broader  than 
long,  apical  margin  narrowly  rounded.  Disc  of  clypeus  weakly 
depressed,  nearly  flat  in  profile  (Fig.  15);  broad  median  line 
impunctate,  punctures  fine  on  either  side  and  mostly  sepa- 
rated by  about  1.5  times  a puncture  diameter,  interspaces 
shiny,  a little  less  so  basad.  Frons  slightly  to  moderately  shiny 
between  fine,  subcontiguous  punctures;  ocellocular  area 
moderately  shiny,  impunctate  adjacent  to  ocellus,  densely  to 
subcontiguously  punctate  near  eye,  with  fine  and  minute 
punctures;  preoccipital  area  slightly  shiny  between  dense  to 
subcontiguous  fine  punctures;  gena  shiny  between  irregularly 
close  to  subcontiguous  punctures  which  are  minute  near  eye, 
becoming  fine  to  moderate  ventrad.  Interantennal  distance 
3.41  (3.13-3.48)  times  antennal  socket  diameter;  antennoc- 
ular  distance  1.63  (1.57-1.63)  times  antennal  socket  diam- 
eter; scape  slender,  2.29  (2.23-2.36)  times  longer  than  wide, 
scape  length  0.72  (0.70-0.75)  times  length  of  first  flagellar 
segment;  first  flagellar  segment  5.28  (5.28-5.50)  times  length 
of  second  and  distinctly  longer  than  combined  length  of  fol- 
lowing three  segments.  Interocellar  distance  2.09  (2.00-2. 1 2) 
times  diameter  of  anterior  ocellus;  ocellocular  distance  1.31 
( 1 .32-1 .45)  times  diameter  of  anterior  ocellus;  ocelloccipital 
distance  1 .88  ( 1 .88-2.03)  times  diameter  of  anterior  ocellus. 

Thorax.  Mesoscutum  shiny  between  fine,  dense  punctures; 
scutellum  similar,  but  punctures  dense  to  subcontiguous  and 
narrow  median  line  impunctate;  metanotum  moderately  shiny 
between  scattered  minute  punctures;  meso-  and  metepisterna 
shiny  between  dense,  fine  punctures.  Basal  area  of  propo- 
deum  moderately  shiny,  sharply  tessellate  and  with  sparse 
fine  punctures  except  along  anterior  margin;  propodeal  side 
shiny  between  dense,  fine  punctures.  Basitibial  plate  (Fig.  3 1 ) 
about  twice  longer  than  wide,  lower  margin  broadly  rounded; 
lower  margin  of  secondary  plate  rounded. 

Abdomen.  First  three  terga  shiny  between  dense,  minute 
punctures;  fourth  and  fifth  terga  shiny  between  dense,  fine 
punctures  which  are  a little  more  separated  than  on  basal 
segments;  pygidial  plate  (Fig.  40)  narrow,  apex  narrowly  trun- 
cate; secondary  plate  with  margins  slightly  concave  toward 
gradually  narrowed  apical  ridge. 

Color.  Head,  thorax,  and  legs  blackish  brown;  scutellum 
dull  reddish;  first  to  fifth  abdominal  terga  metallic  blue-green, 
fourth  and  fifth  more  greenish;  abdominal  sterna  brownish, 
with  obscure  metallic  greenish  tints,  especially  laterad.  The 
following  whitish:  basal  spot  on  mandible;  labrum,  except 
apex  and  along  basal  margin  (sometimes  reduced  to  lateral 


spots);  inverted  T-shaped  clypeal  mark  (may  be  reduced  to 
narrow  median  stripe  and  sublateral,  preapical  spots);  narrow 
supraclypeal  mark  (present  only  in  holotype);  elongate  spot 
on  malar  area;  broad  mark  on  paraocular  area,  ending  nar- 
rowly on  eye  margin  at,  or  slightly  above,  level  of  lower 
margin  of  antennal  socket;  basal  spot  on  pro-  and  mesotibia 
(preapical  protibial  spot  sometimes  present).  Tegula  reddish. 
Wings  dark  brown;  veins  and  stigma  blackish. 

Pilosity.  Hairs  whitish  on  head,  but  with  broad  preoccipital 
band  of  long,  dark  brown  hairs;  ocellar  area  pale-haired,  but 
with  brownish  hairs  on  frons  anterior  to  ocelli.  Hairs  of  tho- 
rax whitish,  but  with  broad  interalar  band  of  dark  brown 
hairs  and  brown  hairs  on  metepisternum  and  side  of  pro- 
podeum.  Hairs  on  front  face  of  first  tergum  pale;  hairs  on 
dorsal  face  of  first,  and  on  second  and  third  terga  appressed, 
simple,  blackish,  very  short;  hairs  of  fourth  and  fifth  terga 
long,  erect,  mostly  plumose,  whitish;  prepygidial  fimbria 
golden  reddish  to  brown.  Hairs  of  legs  mostly  brownish  black, 
but  with  glistening  whitish  hairs  on  coxae,  trochanters,  and 
posterior  margins  of  profemora  and  protibia;  scopal  hairs 
whitish,  becoming  brownish  distad  on  metabasitarsus. 

MALE.  Measurements  (mm).  Head  width  5.79(5.33-5.85); 
head  length  3.79  (3.44-3.79);  wing  length  12.0;  total  length 
14.0 

Head.  1.53  (1.51-1.56)  times  broader  than  long;  occipital 
margin  nearly  flat  in  frontal  view  and  slightly  below  tops  of 
eyes,  ocelli  anterior  to  occipital  margin;  inner  orbits  very 
strongly  convergent  above,  upper  frontal  width  0.80  (0.72- 
0.82)  times  lower  frontal  width.  Mandible  slender,  tridentate, 
inner  tooth  broad  and  with  its  apical  margin  concave.  La- 
brum about  1.6  times  broader  than  long,  moderately  shiny 
between  sparse  fine  punctures  and  with  interspersed  minute 
punctures.  Disc  of  clypeus  nearly  flat  in  profile,  about  as  in 
female,  moderately  shiny,  with  broad  median  impunctate 
line,  punctures  on  either  side  fine,  mostly  separated  by  about 
a puncture  diameter.  Punctation  of  remainder  of  head  about 
as  in  female.  Interantennal  distance  2.72  (2.47-2.81)  times 
diameter  of  antennal  socket;  antennocular  distance  0.79  (0.77- 
0.93)  times  diameter  of  antennal  socket;  scape  stout,  2.18 
(2.07-2.18)  times  longer  than  wide,  0.71  (0.70-0.79)  times 
length  of  first  flagellar  segment;  first  flagellar  segment  5.60 
(5.33-5.60)  times  length  of  second  and  longer  than  following 
three  segments  combined.  Interocellar  distance  1.82  (1.84- 
1 .94)  times  diameter  of  anterior  ocellus;  ocellocular  distance 
0.88  (0.84-0.90)  times  diameter  of  anterior  ocellus;  ocelloc- 
cipital distance  1.76  (2.00-2.06)  times  diameter  of  anterior 
ocellus. 

Thorax.  As  described  for  female,  but  mesoscutal  punctures 
subcontiguous  and  scutellum  slightly  depressed  along  mid- 
line. Metafemur  about  twice  longer  than  thick  (dorsoven- 
trally);  metabasitarsus  about  three  times  longer  than  broad, 
posterior  margin  weakly  curved. 

Abdomen.  As  described  for  female;  apex  of  last  tergite 
broadly  bilobate. 

Terminalia.  Sides  of  distal  process  of  seventh  sternite  (Fig. 
5)  slightly  divergent  distad,  apical  margin  weakly  incised; 
setae  short,  sparse,  simple.  Distal  process  of  eighth  sternite 
(Fig.  6)  weakly  narrowed  subbasally;  setae  long,  numerous. 


Contributions  in  Science,  Number  347 


Sneiling:  American  Centridini  19 


Gonostylus  (Figs.  7,  8)  a little  broadened  distad;  ventral  pro- 
cess broadly  rounded. 

Pilosity.  As  described  for  female,  but  hairs  of  apical  ab- 
dominal sternite  pale  brown;  hairs  of  metatibia  and  meta- 
basitarsus  mostly  dark  brownish,  but  with  some  along  pos- 
teroapical  margins  of  metatibia  golden  brown  and  some  along 
posterior  margin  of  metabasitarsus  light  brown,  with  pale 
tips. 

Color.  Body  color  as  described  for  female,  except  whitish 
marks  as  follows:  spot  at  base  of  mandible;  entire  labrum; 
clypeus,  except  along  laterobasal  margin;  broad  supraclypeal 
mark;  spot  on  malar  area;  paraocular  area,  ending  narrowly 
on  inner  eye  margin  slightly  above  level  of  lower  margin  of 
antennal  socket;  underside  of  scape;  basal  spot  on  all  tibiae; 
outer  stripe  on  protibia;  sublateral  mark  at  base  of  second 
tergite.  Tarsi  reddish  brown  to  brown.  Wings  light  brown. 

TYPE  MATERIAL 

Holotype  female  and  allotype:  airstrip,  Isla  El  Rey,  Islas  Per- 
las,  PANAMA,  22  Feb.  1981  (D.W.  Roubik),  flying  around 
Dioclea  megacarpa,  in  Natural  History  Museum  of  Los  An- 
geles County.  Paratypes:  12,  same  data  as  holotype;  5<3<3,  Coi- 
ba  Island,  Veraguas,  PANAMA,  21-24  Oct.  1979  (D.  Rou- 
bik); 12,  Fort  Kobbe,  Canal  Zone,  PANAMA,  9 Sept.  1958 
(W.J.  Hanson);  1 <3,  Patilla  Point,  Canal  Zone,  PANAMA,  15 
Jan.  1929  (C.H.  Curran);  1<3,  Bruja  Point,  Canal  Zone,  PAN- 
AMA, 25  Jan.  1929  (C.H.  Curran);  12,  LaChorrera,  Panama, 
PANAMA,  22  May  1912  (A.  Busck);  1 <5,  5 km  E Comarca 
de  San  Bias,  Puerto  Obaldia,  PANAMA,  1 1 Oct.  1979  (D. 
Roubik);  322,  Tamarindo  Beach,  Guanacaste,  COSTA  RICA, 
28  Feb.  1980  (G.W.  Frankie);  222,  same  locality  and  collec- 
tor, 30  Mar.  1 980,  on  Haematoxylon  brasiletto\  266,  La  Union, 
Playa  El  Icacal,  EL  SALVADOR,  8 July  1975  (E.M.  and  J.L. 
Fisher).  Paratypes  in  AMNH,  LACM,  ROUB,  UCB,  UKAN, 
and  USNM. 

ETYMOLOGY 

Combines  the  Greek  words  aithiops  (swart  or  dark)  and  kes- 
tos  (girdle  or  band). 

DISCUSSION 

Although  this  species  superficially  looks  much  like  C.  flavi- 
frons  and  C.  flavofasciata,  it  is  smaller  than  either.  The  disc 
of  the  clypeus,  in  profile,  is  distinctly  flatter  in  C.  aethiocesta 
than  in  the  other  two  species  (compare  Fig.  1 5 with  Fig.  16) 
and,  in  both  sexes,  the  disc  is  more  closely  and  abundantly 
punctate.  Small  females  of  C.  flavofasciata  may  be  no  larger 
than  females  of  C.  aethiocesta  but  have  the  tibial  scopa  black 
and  the  hairs  of  the  side  of  the  propodeum  are  whitish.  The 
much  larger  size  (head  width  greater  than  7.0  mm)  will  sep- 
arate C.  flavifrons  females  from  C.  aethiocesta,  as  will  the 
sparser  clypeal  punctation  and  the  differently  shaped  basi- 
tibial  plate.  Males  of  C.  aethiocesta  have  the  apical  abdom- 
inal segments  dark  and  immaculate;  in  C.  flavifrons  and  C. 
flavofasciata  males  the  apical  segments  are  usually  reddish 
and  commonly  are  yellowish  maculate;  in  these  two  species, 
the  hairs  of  the  propodeum  are  usually  whitish. 


Two  South  American  species  also  resemble  C.  aethiocesta 
and  are  of  about  the  same  size.  Females  of  C.  buchwaldi 
Friese,  1 900,  have  a dark  scopa,  pale  propodeal  hairs,  a band 
of  prostrate,  simple,  golden  brown  hairs  across  the  apical 
margin  of  the  third  tergite,  and  the  erect  hairs  of  the  fourth 
tergite  are  simple.  In  females  of  C.  nigrofasciata  Friese,  1 899, 
there  is  a band  of  pale  hairs  across  the  apical  margin  of  the 
third  tergite,  the  apical  margin  of  the  fourth  tergite  is  fer- 
ruginous, the  fifth  tergite  is  ferruginous  and  with  simple  erect 
hairs,  and  the  basitibial  plate  is  more  elongate,  with  a more 
acute  apex. 

In  the  males  of  both  C.  buchwaldi  and  C.  nigrofasciata  the 
clypeus  is  only  slightly  shiny,  with  the  integument  conspic- 
uously roughened.  Neither  of  these  has  brown  hairs  on  the 
side  of  the  propodeum  and  in  both  species  the  fifth  and  sixth 
tergites  are  ferruginous.  Males  of  C.  nigrofasciata  also  differ 
in  that  the  ocellocular  distance  is  greater  than  the  diameter 
of  the  anterior  ocellus. 

Centris  ( Centris ) flavifrons  (Fabricius) 

Figures  14,  29,  38 

Apis  flavifrons  Fabricius,  1775:383.  <5. 

Apis  flavifrons  brasiliana  Christ,  1791:1 40. 

Centris  flavifrons:  Lepeletier,  1841:152.  F.  Smith,  1874:361. 
Centris  citrotaeniata  Gribodo,  1894:267.  2.  NEW  SYN- 
ONYMY. 

Centris  flavifrons  var.  nigritula  Friese,  1899:46.  2.  NEW 
SYNONYMY. 

Centris  flavifrons  var.  rufescens  Friese,  1899:46.  <3.  NEW 
SYNONYMY. 

Centris  (Cyanocentris)  flavifrons:  Friese,  1 900b:3 1 7—3 1 8 (in 
part)  (tax.,  distr.). 

Centris  ( Centris ) flavifrons:  Michener,  1954:137  (distr.). 

Moure,  1 960b:  1 25-126  (tax.). 

Centris  (Centris)  rufescens:  Michener,  1 954: 1 37  (tax.,  status). 
“Centris  fulvifrons"  Moure,  1960a:  10  (lapsus  for  C.  flavi- 
frons). 

Centris  ( Centris ) citrotaeniata:  Moure,  1960a:  10-1 1 (tax.). 
Centris  (Centris)  nigritula:  Snelling,  1974:30-31  (tax.  status). 

Moure  (1960b)  examined  and  redescribed  the  type  of  C. 
flavifrons,  from  “Brasilia.”  He  noted  that,  according  to  cur- 
rent concepts,  C.  flavifrons  ranges  from  southern  Brazil  to 
Mexico.  Throughout  this  range  this  is  often  a common  bee 
and  one  subject  to  considerable  variation.  It  is  now  apparent 
that  my  effort  to  separate  C.  f.  nigritula  as  a distinct  species 
(Snelling,  1974)  was  futile.  Since  that  time  I have  examined 
several  hundred  specimens  and  seen  so  broad  a range  of 
variation  that  it  is  now  apparent  that  C.  nigritula  is  not 
worthy  of  recognition  at  any  level. 

Friese  (1899)  described  C.  flavifrons  var.  rufescens  from  a 
male  from  Chiriqui,  Panama.  It  was  said  to  differ  from  the 
nominate  form  in  possessing  wide,  yellow  abdominal  bands 
and  reddish  pubescence  on  the  legs.  Although  Michener  (1954) 
thought  that  this  seemed  to  represent  a distinct  species,  I do 
not  agree.  Neither  Michener  nor  I has  seen  the  type  of  var. 
rufescens  but  I suspect  that  it  is  nothing  more  than  a metan- 
der.  Metanders  of  both  C.  flavifrons  and  C.  flavofasciata  are 


20  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


characterized  by  possessing  broad  abdominal  fasciae  and  fer- 
ruginous pubescence  on  the  legs.  Since  nothing  in  Friese’s 
brief  description  would  indicate  to  which  of  these  species  this 
form  should  be  assigned,  I have  arbitrarily  elected  to  follow 
Friese  and  include  it  under  C.  flavifrons.  as  a synonym.  In 
Panama,  C.  flavifrons  is  more  common  than  is  C.  flavofas- 
ciata  and  it  seems  more  likely  than  the  var.  rufescens  belongs 
with  the  former  species. 

Centris  citrotaeniata  was  also  described  from  Chiriqui, 
Panama,  based  on  a single  female  specimen.  The  type  is  in 
the  collections  of  the  Museu  Civico  de  Storia  Naturale,  Gen- 
oa, Italy,  and  was  examined  by  Moure  ( 1 960a).  Moure  noted 
that  this  species  was  close  to  “C.  fulvifrons evidently  a 
lapsus  for  C.  flavifrons.  From  the  original  description,  as  well 
as  the  supplemental  notes  provided  by  Moure,  it  is  clear  that 
C.  citrotaeniata  is  equivalent  to  dark  Panamanian  individ- 
uals of  C.  flavifrons.  and  so  Gribodo’s  species  is  here  placed 
in  synonymy. 

Variation  in  females  of  C.  flavifrons  chiefly  involves  the 
extent  of  pale  versus  blackish  pubescence  on  the  thorax  and 
on  the  scopa.  In  females  from  Brazil  the  thoracic  hairs  are 
mostly  whitish,  with  a conspicuous  broad  interalar  band  of 
blackish  pubescence;  a few  dark  brown  hairs  are  present  be- 
low the  wing  bases  and  the  scopa  is  uniformly  pale.  In  ma- 
teria! from  Guyana  and  French  Guiana,  there  is  considerable 
replacement  of  the  pale  pleural  hairs  by  blackish  hairs  and 
the  scopa  is  largely  dark,  but  with  conspicuous  white  hairs 
anteriorly  on  both  the  tibia  and  basitarsus.  This  dark  variant 
also  has  many  dark  hairs  on  the  fourth  and  fifth  terga.  A 
similar  variant  also  appears  sporadically  in  samples  from 
Panama  and  Costa  Rica.  The  darkest  phenotype  is  from 
Trinidad,  the  var.  nigritula.  In  these  specimens,  the  thorax 
is  entirely  blackish  pubescent,  with  only  the  axilla,  scutellum, 
and  thoracic  venter  whitish  pubescent.  Curiously,  the  scopa 
is  predominantly  pale,  with  only  a few  brownish  hairs  distally 
on  the  tibia  and  along  the  posterior  margin  of  the  basitarsus. 
Some  of  the  Panamanian  females  have  the  thoracic  hairs 
wholly  pale,  except  for  a small  median  patch  of  dark  hairs 
on  the  mesoscutum;  in  these  individuals  the  scopa  is  mainly 
dark. 

The  Central  American  populations  vary  on  a smaller  scale. 
Specimens  from  Mexico  typically  have  a large  inverted 
T-shaped  clypeal  mark,  the  labrum  is  mostly  dark  and  the 
lateral  face  mark  fills  the  paraocular  area  below  the  level  of 
the  tentorial  pit.  The  transverse  mesoscutal  fascia  of  dark 
hair  is  preceded  by  a narrow  band  of  white  pubescence,  the 
mesopleuron  is  largely  dark  pubescent,  and  the  tibial  scopa 
is  only  about  one-half  pale.  Coloration  becomes  progressively 
darker  toward  the  south  and  specimens  from  Costa  Rica  may 
be  much  darker:  the  labrum  wholly  black,  the  clypeal  mark 
broken  into  two  or  three  narrow  segments,  and  the  meso- 
scutum with  only  a small  cluster  of  pale  hairs  on  the  antero- 
median portion;  the  scopa  is  almost  entirely  dark,  with  pale 
hairs  anteriorly  on  the  metatibia  and  anterobasally  on  the 
metabasitarsus.  Another  characteristic  of  the  Costa  Rican 
specimens  is  that  the  hairs  of  the  dorsal  face  of  the  first  tergite 
are  blackish.  In  the  Mexican  specimens  the  hairs  are  pale  at 
the  sides  and  across  the  basal  portion  of  the  dorsal  face. 


The  tendency  toward  increasing  melanism  is  partially  re- 
versed in  Panama.  A long  series  from  Panama  Province, 
collected  by  R.W.  Brooks,  exhibits  considerable  variation. 
In  some  specimens  the  interalar  fascia  is  nearly  absent  and 
in  others  it  may  cover  up  to  two-thirds  of  the  mesoscutum. 
In  most  specimens  the  mesepisternum  has  only  a few  dark 
hairs  below  the  tegula  and  in  others  there  may  be  a blackish 
hair  patch  over  the  upper  one-half  or  more  of  the  segment. 
The  scopa  varies  from  nearly  wholly  pale  to  about  one-half 
black.  Face  marks  may  be  greatly  reduced:  mandible  and 
labrum  entirely  dark;  clypeus  with  narrow  median  line  and 
widely  separated  apicolateral  spots  and  scape  wholly  dark. 
Reduction  of  face  marks  is  not  correlated  with  increased 
melanism  in  pubescence.  Many  of  these  specimens  will  match 
the  color  pattern  of  the  type  of  C.  citrotaeniata. 

The  darkest  Costa  Rican  specimens  superficially  resemble 
the  form  from  Guyana,  French  Guiana,  and  Trinidad  de- 
scribed by  Friese  as  the  var.  nigritula.  They  differ  from  that 
form,  however,  in  that  the  plumose  hairs  of  the  fourth  tergite 
are  uniformly  pale.  In  the  var.  nigritula  there  are  many  dark 
hairs  on  the  fourth  tergite  and,  in  some  specimens,  pale  hairs 
may  be  entirely  lacking. 

In  general,  then,  this  species  becomes  progressively  more 
melanic,  proceeding  from  central  Brazil  to  northeastern  South 
America.  Westward  across  northern  South  America  there  is 
a reversal,  toward  decreased  melanism,  in  Venezuela,  Co- 
lombia, and  Panama.  The  melanic  trend  is  again  evident  in 
Costa  Rica  and  Nicaragua,  but  decreases  once  more  at  the 
northern  (Mexican)  extremities  of  the  range.  There  are  nu- 
merous localized  phenotypes  which  may  depart,  in  one  way 
or  another,  from  the  general  dine. 

The  discussion  thus  far  has  focused  primarily  on  the  fe- 
males, though  the  pubescent  patterns  of  the  males  of  C.  flavi- 
frons generally  follow  those  of  the  females.  There  are,  how- 
ever, problems  that  are  peculiar  to  the  males.  Both  this  species 
and  the  similar  C.  flavofasciata  possess  metanders,  males 
which  are  unusually  large  and  robust  and  much  more  exten- 
sively marked  with  bright  yellow,  especially  on  the  legs  and 
abdomen.  While  it  is  possible  to  separate  normal  males  of 
the  two  species,  the  metanders  are  much  less  convincingly 
handled. 

In  part,  this  is  due  to  a scarcity  of  fully  developed  metan- 
ders which  can  be  assigned  with  surety  to  one  or  the  other 
of  these  two  species.  And,  in  part,  the  metanders  that  are 
available  tend  not  to  exhibit  the  differences  which  separate 
the  normal  males.  I do  not  believe  that  this  in  any  way 
invalidates  the  distinctions  between  the  two  species,  but  it 
is  merely  another  manifestation  of  a common  problem  within 
some  subgenera:  within  a given  complex  males  may  be  much 
less  readily  separated  than  their  respective  females.  A final 
difficulty  is  that,  unlike  the  situation  in  such  subgenera  as 
Xerocentris  and  Paracentris,  the  metandrous  and  normal 
males  form  a more  or  less  continuous  series,  with  many 
individuals  of  intermediate  stature. 

A series  of  about  30  males  collected  by  J.A.  Chemsak  near 
Mazatlan,  Sinaloa,  Mexico,  12  Dec.  1980  (UCB)  consists 
largely  of  metanders  of  C.  flavofasciata.  but  with  a few  nor- 
mal males  and  intermediates.  While  I have  seen  many  males 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  21 


of  C.  flavifrons  that  seem  to  be  intermediates,  I have  only 
three  that  are  clearly  fully  developed  metanders,  two  from 
Brazil  and  one  from  Mexico. 

Normal  males  of  C.  flavifrons  possess  the  following  char- 
acteristics, each  subject  to  moderate  variation:  the  lower  in- 
ner mandibular  carina  terminates  in  a low,  obtusely  trian- 
gular process  (Fig.  14);  the  clypeal  disc  is  smooth  and  polished 
throughout  between  fine,  scattered  punctures  and  is  not  whol- 
ly yellow;  the  laterobasal  black  mark  of  the  clypeus  extends 
onto  the  disc  and  sometimes  the  clypeus  bears  only  an  in- 
verted (though  broad)  T-mark.  The  second  to  fourth  abdom- 
inal terga  are  maculate;  the  maculation  of  the  second  segment 
often  consists  merely  of  two  somewhat  elongate  sublateral 
spots,  but  may  extend  across  the  base  of  the  segment  as  a 
continuous  or  narrowly  interrupted  band;  when  it  is  a band, 
it  is  almost  always  much  shorter  in  the  middle  than  the  dark 
area  distad  to  it. 

In  normal  males  of  C.  flavofasciata  the  lower  inner  man- 
dibular carina  ends  in  an  elongate,  acute,  tooth-like  process. 
The  disc  of  the  clypeus  is  wholly  yellow  and  the  areas  on 
either  side  of  the  shiny,  impunctate  median  line  are  less  shiny 
and  distinctly  roughened  between  fine  punctures  that  are 
mostly  separated  by  1.0  to  1.5  times  a puncture  diameter. 
The  abdominal  terga  are  almost  always  continuously  fasciate 
and  the  yellow  band  of  the  second  segment  is  as  long  as,  or 
longer  than,  the  dark  area  that  follows  it. 

Most  of  these  distinctions  disappear  in  the  metanders,  or 
are  at  least  less  certain.  Some  metanders  of  C.  flavofasciata 
have  the  clypeal  disc  nearly  polished  and  the  punctures  about 
as  sparsely  distributed  as  in  C.  flavifrons.  While  very  nearly 
all  metanders  of  C.  flavofasciata  have  the  clypeal  disc  wholly 
yellow,  there  are  a few  in  which  the  laterobasal  black  mark 
does  extend  slightly  onto  the  disc.  The  abdominal  banding 
breaks  down  completely,  since  metanders  of  both  species 
have  broad,  continuous  yellow  fasciae  on  the  second  to  sixth 
segments  and  an  interrupted  band  across  the  dorsum  of  the 
first  segment. 

The  only  feature  that  seems  to  hold  up  is  the  shape  of  the 
termination  of  the  lower  inner  mandibular  carina.  In  me- 
tanders of  C . flavifrons  the  terminus  is  low  and  obtuse,  much 
as  in  the  normal  males.  It  is  an  acutely  tooth-like  process  in 
C.  flavofasciata.  How  consistent  this  difference  is,  I cannot 
now  state,  only  that  it  does  hold  true  for  the  specimens  ex- 
amined thus  far. 

There  is  obviously  much  to  be  done  before  the  phenom- 
enon of  metandry  can  be  adequately  dealt  with  taxonomi- 
cally.  Presumably,  the  metanders  of  such  species  as  C.  flavi- 
frons and  C.  flavofasciata  are  behaviorally  different  from 
their  normal  counterparts,  perhaps  in  a manner  similar  to 
the  situation  in  such  species  as  C.  (Xerocentris)  pallida  W. 
Fox,  the  only  species  studied  thus  far  (Alcock  et  al.,  1976, 
1977).  In  that  species  metanders  seek  newly  emerging  females 
at  nest  sites  of  the  previous  year.  The  normal  males  establish 
territories  at  nectar/pollen  sources  and  attempt  to  mate  with 
females  there. 

A large  nesting  aggregation  of  C.  flavifrons  was  observed 
in  Panama.  Normal  males  and  the  rufescent  variety  were 
present.  Males  were  observed  to  “.  . . purposely  attack  Me- 


soplia  in  the  nest  area— with  a frequency  far  higher  than  the 
pursuit  of  other  males  or  females”  (D.  Roubik  and  D.  Yane- 
ga,  in  prep.). 

Centris  (Centris)  flavofasciata  Friese 

Figures  16,  30,  39 

Centris  flavifrons  var.  flavofasciata  Friese,  1899:46.  <5. 
Centris  ( Cyanocentris ) flavifrons  var.  flavofasciata:  Friese, 

1900b:318.  $ (distr. , tax.). 

Centris  (Centris)  flavofasciata:  Michener,  1954:137  (distr.). 

Snelling,  1966:23  (distr.). 

Centris  flavofasciata  is  a common  Mexican  species  which 
ranges  from  southern  Arizona  to  Panama.  It  is  possible  that 
the  range  extends  into  northern  South  America,  but  I have 
seen  no  specimens  to  substantiate  that  assumption.  Most  of 
the  features  by  which  this  species  may  be  separated  from 
other  members  of  the  C.  flavifrons  groups  are  summarized 
in  the  keys. 

Females  of  C.  flavofasciata  consistently  have  a dark  scopa, 
with  a few  of  the  posteroapical  hairs  of  the  metabasitarsus 
with  pale  tips.  Most  females  have  a small  patch  of  dark  hairs 
below  the  base  of  the  forewing;  none  has  the  side  of  the  thorax 
extensively  dark  pubescent,  as  is  usually  true  of  Central 
American  material  of  C.  flavifrons.  Those  specimens  of  C. 
flavifrons  with  little  dark  pleural  pubescence  are  from  Pan- 
ama. 

Facial  maculations  of  C.  flavofasciata  tend  to  be  somewhat 
whitish  in  females,  rather  than  definitely  yellow  as  in  C. 
flavifrons  and,  in  both  sexes,  are  more  extensive;  in  the  male 
the  entire  clypeal  disc  is  yellow,  not  encroached  upon  by  the 
black  laterobasal  mark.  In  females  of  C.  flavofasciata  the 
labrum  is  largely  pale,  rather  than  extensively  dark  and  the 
vertical  and  lateral  arms  of  the  inverted  T-mark  of  the  clyp- 
eus are  broad  and  never  broken  into  segments. 

Ptilocentris,  new  subgenus 

DIAGNOSIS 

Differs  from  other  subgenera  of  Centris  by  the  following  com- 
bination of  characters.  Maxillary  palp  five-segmented;  tergal 
integument  metallic  blue-green,  first  four  terga  with  pubes- 
cence long,  dense,  erect,  plumose.  Female:  secondary  basi- 
tibial  plate  with  distinct  overhanging  margin,  pro-  and  me- 
sobasitarsi  with  elaiospathe;  clypeal  disc  evenly  rounded  into 
lateral  and  basal  faces,  gently  convex  from  side  to  side.  Male: 
metabasitarsus  without  posterior  carina;  genitalia  without 
branched  setae;  upper  inner  mandibular  carina  ending  at  base 
of  inner  tooth;  eighth  sternite  expanded  at  apex. 

DESCRIPTION 

Mandible  narrow,  neither  subapically  broadened  nor  bent; 
tri-  (male)  or  quadridentate  (female);  upper  inner  carina  end- 
ing near  base  of  innermost  tooth;  lower  inner  carina  not 
subangularly  produced,  continuous  to  innermost  tooth  along 
upper  margin.  Labrum  of  female  about  1.3  times  broader 
than  long,  apex  subacute;  about  as  broad  as  long  in  male. 


22  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


Clypeus  broader  than  long  and  separated  from  inner  eye 
margin  by  less  than  diameter  of  anterior  ocellus;  disc  not  at 
all  flattened,  low-convex  from  side  to  side  and  evenly  round- 
ed onto  lateral  faces  and  gently  sloping  toward  base.  First 
flagellar  segment  about  as  long  as  following  three  combined 
(female)  or  shorter  (male).  Ocellocular  distance  greater  than 
anterior  ocellus  diameter.  Pro-  and  mesobasitarsi  with  an- 
terior elaiospathe  on  inner  surface;  basitibial  plate  of  female 
with  distinct  secondary  plate,  its  lower  margin  overhanging 
primary  plate;  metafemur  of  male  robust,  without  ventral 
ridge  or  tubercle;  metabasitarsus  simple,  without  posterior 
ridge  or  carina.  Female  pygidial  plate  V-shapcd,  apex  nar- 
rowly rounded;  secondary  plate  distinct,  apex  acute.  Male 
without  distinct  pygidial  plate;  seventh  sternite  (Fig.  9)  sub- 
apically  broadened  and  apical  margin  deeply  emarginate;  with 
abundant  long,  plumose  hairs  at  apex,  with  short,  simple 
hairs  at  base  of  apical  lobe;  eighth  sternite  (Fig.  10)  with 
distal  process  long,  its  apex  angularly  spatulate,  apical  one- 
half  of  shaft  and  broadened  distal  portion  with  long,  plumose 
hairs;  genital  capsule  (Figs.  11,  12)  robust,  gonocoxite  mas- 
sive; gonostylus  short,  broad,  with  a few  fine,  simple  setae. 
Abdominal  terga  metallic  blue-green  and  four  basal  segments 
with  abundant,  erect,  plumose  hairs  on  discs. 

TYPE-SPECIES 

Centris  festiva  F.  Smith,  1854,  by  present  designation. 

ETYMOLOGY 

The  subgeneric  name  combines  Greek  ptilon  (feather)  with 
the  generic  name  Centris  and  refers  to  the  presence  of  abun- 
dant long,  erect,  plumose  hairs  on  the  abdominal  terga. 

The  subgeneric  name  proposed  here  was  originally  con- 
ceived by  J.S.  Moure  and  appears  on  specimens  of  C.  festiva 
in  various  collections  which  he  identified  20  years  ago.  It  is 
an  appropriate  name  and  I am  pleased  to  adopt  it  here. 

DISCUSSION 

The  type-species  is  the  only  known  representative  of  this 
subgenus. 

Ptilocentris  seems  to  be  another  segregate  of  the  Paracen- 
tris-Centris  s.  str.  complex  of  subgenera.  In  my  key  to  the 
subgenera  of  Centris  (Snelling,  1974)  the  female  will  go  to 
Centris  s.  str. 

Females  differ  from  those  of  the  nominate  subgenus  in 
lacking  a stipital  comb;  in  Centris  s.  str.  the  comb  consists 
of  a long  row  of  stout,  close-set,  coarse,  acute  bristles.  The 
labrum  in  Centris  s.  str.  is  much  broader,  with  the  apical 
margin  broadly  rounded.  The  broad,  distinctly  flattened,  or 
slightly  depressed,  clypeal  disc  which  abruptly  slopes  basad 
and  toward  the  lateral  margins  is  also  distinctive  of  Centris 
s.  str.  In  this  regard,  Ptilocentris  is  more  like  some  species 
of  Paracentris  from  which  it  immediately  differs  in  possessing 
a sharply  defined  secondary  basitibial  plate  with  a distinct 
margin  which  extends  slightly  over  the  disc  of  the  primary 
plate.  Also,  Paracentris  is  like  Centris  s.  str.  in  the  form  of 
the  comb  of  the  stipes. 


The  male  of  Ptilocentris  is  unique  among  the  Paracentris - 
Centris  s.  str.  complex  in  the  form  of  the  gemlalic  structures. 
The  shapes  of  the  seventh  and  eighth  stermtes  are  distinctive 
and  the  lack  of  coarse,  plumose  setae  on  the  genitalic  capsule 
is  also  an  unusual  feature  in  this  complex.  So,  too,  is  the 
short,  broad  gonostylus. 

Centris  (Ptilocentris)  festiva  F.  Smith 

Figures  9-12 

Centris  festiva  F.  Smith,  1854:375.  2. 

Centris  chlorura  Cockerell,  1919:188-189.  2.  NEW  SYN- 
ONYMY. 

This  attractive  and  unusual  species  is  easily  recognized  by 
the  characteristic  pattern  of  erect  hairs  on  the  discs  of  the 
abdominal  terga.  In  most  specimens  the  second  and  third 
terga  have  a basal  zone  of  blackish  hairs,  but  in  some  spec- 
imens the  hairs  may  be  wholly  pale;  one  such  pale  haired 
specimen  was  the  basis  for  C.  chlorura,  the  type  of  which 
has  been  examined. 

There  are  no  previous  records  of  this  species  from  Central 
America.  In  South  America,  C.  festiva  is  known  to  be  present 
in  Colombia,  Ecuador,  and  Peru. 

NEW  RECORDS 

MEXICO,  DISTRITO  FEDERAL:  13,  Los  Venados,  6 Nov. 
1938  (G.  Vivas-Berthier;  USNM).  COSTA  RICA,  ALAJUE- 
LA:  1<3,  5 km  S Vara  Blanca,  1 1 Nov.  1973  (P.A.  Opler; 
LACM).  SAN  JOSE:  12,  San  Jose,  “1.6.19”  (M.  Valerio; 
USNM).  PANAMA,  CHIRIQUI:  13,  Osra  Clara,  26  Jan. 
1981  (D.W.  Inouye;  ROUB).  According  to  Roubik  (pers. 
comm.):  “Sandy  Knapp  also  collected  C.  festiva  in  Chinqui 
on  22  April  82  ...  at  Cerro  Colorado,  about  1000  m ele- 
vation.” 

Subgenus  Melanocentris  Friese 

Centris  subg.  Melanocentris  Friese,  1900b:241,  244.  Type- 
species:  Centris  atra  Friese,  1900b;  designated  by  Sand- 
house,  1943. 

Species  belonging  to  the  subgenus  Melanocentris  are  medi- 
um-sized to  large  bees,  usually  black  (though  with  pale  face 
marks  in  the  males)  or  with  the  abdomen  more  or  less  dusky 
ferruginous.  Less  commonly  the  abdomen  may  have  metallic 
blue  or  green  reflections.  The  pubescence  is  mostly  dark  but 
there  may  be  limited  amounts  of  pale  pubescence  on  the 
thorax,  the  hind  legs,  and  on  the  abdomen.  In  both  sexes  the 
maxillary  palp  is  five-segmented  and  the  mandible  is  stout, 
quadridentate  in  the  female  and  tndentate  in  the  male;  in 
the  female  the  mandible  is  broadened  and  abruptly  bent  near 
the  apex. 

Additional  characteristics  of  the  female  include  the  pres- 
ence of  an  elaiospathe  on  the  pro-  and  mesobasitarsi,  the 
weakly  bilobate  scutellum,  the  upper  inner  carina  of  the  man- 
dible ends  near  the  base  of  the  subbasal  tooth,  and  the  margin 
of  the  secondary  basitibial  plate  overhangs  the  primary  plate. 
Males  lack  giant  branched  setae  on  the  genitalia,  the  meta- 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  23 


11  12 


Figures  9-12.  Centris  (Plilocentris)  festiva,  male  seventh  and  eighth  sternites  and  genitalia  (ventral  and  dorsal  views).  Scale  line  = 1.00  mm. 


24  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


Figures  13-14,  apex  of  left  mandible  of  male;  13,  Centris  (C.)  aethiocesta\  14,  C.  (C.)  flavifrons.  Figs.  15-16,  clypeal  profile  of  female:  15,  C. 
(C.)  aethiocesta\  16,  C.  (C.)  Jlavofasciata.  Fig.  17,  apex  of  left  mandible  of  male  C.  (Melanocentris)  fusciventris.  Figs.  18-19,  melabasitarsus 
of  male:  18,  C.  ( Trachina ) labiata\  19,  C.  (T.)  longimana. 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  25 


basitarsus  is  without  an  elevated  carina  on  the  posterior  mar- 
gin, the  scutellum  is  weakly  swollen  on  either  side  of  the 
middle,  and  the  upper  inner  mandibular  carina  ends  near  the 
base  of  the  middle  tooth. 

This  is  primarily  a South  American  group,  and  much  in 
need  of  revisionary  study.  The  few  North  American  species 
may  be  separated  by  the  following  key. 

KEY  TO  NORTH  AMERICAN  MELANOCENTRIS 

1 a.  Male,  antenna  1 3-segmented  and  basiti bial  plate  absent 

2 

b.  Female,  antenna  12-segmented  and  basitibial  plate 

present  8 

2a.  Integument  of  abdominal  terga  shiny,  reddish  or  black- 
ish, never  strongly  metallic;  if  dull,  punctures  subcon- 
tiguous  to  dense,  sharply  defined  at  least  on  second 
tergunr,  metafemur  swollen,  but  without  ventral  ridge 

3 

b.  Integument  of  abdominal  terga  dull  to  slightly  shiny, 
dark  blue,  punctures  on  discs  of  second  and  third  ter- 
gites  very  fine,  obscured  by  dense  tessellation  and  well 
separated;  metafemur  swollen,  with  low  ventral  ridge 

which  terminates  slightly  beyond  middle 

plumipes  F.  Smith 

3a.  Pilosity  of  thorax  buff-colored  to  ferruginous  (hairs  of 

mesoscutum  may  be  dark-tipped)  4 

b.  Pilosity  of  thorax  dark  brown  to  blackish  (one  species 
with  light  hairs  on  scutellum,  metanotum,  and  pro- 

podeum)  5 

4a.  Inner  mandibular  tooth  broad,  cutting  margin  incised 
so  that  mandible  approaches  quadridentate  condition 
(Fig.  1 7);  pilosity  of  third  and  following  tergites  blackish 

fusciventris  Mocsary 

b.  Inner  mandibular  tooth  narrow,  triangular,  mandible 
definitely  tridentate;  pilosity  of  third  and  following  ter- 
gites ferruginous flavilabris  Mocsary 

5a.  Clypeus  narrow,  about  1 . 10  to  1 .25  times  broader  than 
long;  lower  facial  width  no  more  than  1 .5  times  broader 
than  interocular  distance,  latter  always  greater  than 

transocellar  distance  6 

b.  Clypeus  broader,  at  least  1.40  times  broader  than  long; 
lower  facial  width  at  least  1.6  (and  usually  more  than 
1 .8)  times  minimum  interocular  distance,  latter  usually 

no  more  than  transocellar  distance,  often  less 7 

6a.  Abdomen  dusky  ferruginous;  posterior  part  of  thorax, 
metatibia  and  metabasitarsus  pale  pilose;  disc  of  second 
tergum  dull,  punctures  mostly  separated  by  less  than  a 

puncture  diameter agiloides,  new  species 

b.  Abdomen  brownish  to  blackish;  pilosity  of  thorax  and 
leggs  blackish;  disc  of  second  tergum  moderately  shiny 
between  punctures  mostly  separated  by  a puncture  di- 
ameter or  more  sericea  Friese 

7a.  Dorsal  face  of  scutellum  depressed  along  midline  and 
slightly  raised  on  either  side,  apex  of  eminence  im- 
punctate  or  nearly  so;  basal  area  of  propodeum  slightly 
shiny  and  conspicuously  tessellate;  pubescence  of  legs 
dark  obsoleta  Lepeletier 


b.  Dorsal  face  of  scutellum  neither  depressed  along  mid- 
line nor  raised  on  either  side,  uniformly  subcontig- 
uously  punctate;  basal  area  of  propodeum  shiny,  with 
very  weak  tessellation;  hairs  pale  on  metatibia  and  me- 
tabasitarsus   agilis  F.  Smith 

8a.  Apex  of  pygidial  plate  broad,  deeply  and  angularly  in- 
cised (Fig.  42);  scopa  pale;  yellowish  marks  often  pres- 
ent on  some  part  of  lower  face  9 

b.  Apex  of  pygidial  plate  narrowly  truncate  or  acute,  never 
incised  (Figs.  41,43);  scopa  often  dark;  face  rarely  mac- 
ulate   10 

9a.  First  four  abdominal  terga  black,  with  strong  metallic 

blue  reflections species  A 

b.  Entire  abdomen  dusky  ferruginous  

flavilabris  Mocsary 

10a.  Lower  half  of  clypeal  disc  slightly  shiny  to  shiny,  but 
conspicuously  roughened  or  ridged  between  punctures; 
labrum  and  side  of  clypeus  black;  paraocular  area  usu- 
ally black;  integument  of  frons  and  mesopleuron  black, 
without  metallic  bluish  or  greenish-bronze  reflections 

11 

b.  Lower  half  of  clypeal  disc  smooth  and  polished  between 
punctures;  most  of  labrum,  at  least  side  of  clypeus,  and 
paraocular  area  yellowish  maculate;  frons  and  meso- 
pleuron with  metallic  bluish  or  greenish-bronze  reflec- 
tions   plumipes  F.  Smith 

1 la.  Discs  of  second  and  third  terga  moderately  to  strongly 
shiny,  punctures  various  but  usually  not  subcontiguous 
and  much  coarser  than  hairs  arising  from  them  (if  sub- 
contiguous,  scopa  dark);  scopa  often  dark;  abdomen 

often  black  12 

b.  Discs  of  second  and  third  terga  dull,  contiguously  punc- 
tate, punctures  much  coarser  than  hairs  arising  from 

them;  scopa  pale;  abdomen  ferruginous  

agiloides,  new  species 

12a.  Punctures  of  disc  of  second  tergum,  when  visible,  little 
larger  than  hairs  arising  from  them  and  finer  than  those 
on  disc  of  following  segment;  pubescence  of  thorax  and/ 
or  scopa  often  partly  pale;  basitibial  plate  and  pygidium 

various 13 

b.  Punctures  of  disc  of  second  tergum  subcontiguous,  much 
coarser  than  hairs  arising  from  them  and  conspicuously 
coarser  than  punctures  of  apical  zone  and  of  following 
segment;  thoracic  and  scopal  hairs  black;  secondary 
basitibial  plate  acute  (Fig.  24);  pygidial  plate  narrowly 

V-shaped,  secondary  plate  short  and  broad 

sericea  Friese 

13a.  Dorsal  face  of  first  tergum,  across  middle  one-third, 
polished,  without  evident  punctures  and  with  only  a 
few  widely  scattered,  appressed  simple  hairs;  pygidial 
plate  broadly  V-shaped  and  broadly  truncate  at  apex, 
lateral  margins  somewhat  convex  at  about  midlength, 
secondary  plate  elongate  and  with  median  raised  ridge 
extending  from  its  apex  to  apical  truncation  (not  always 

visible  in  worn  specimens)  (Fig.  43);  scopa  pale 

agilis  F.  Smith 

b.  Dorsal  face  of  first  tergum  pubescent  and/or  punctate 
across  middle  half  at  least  at  summit  of  declivity;  py- 


26  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


gidium  not  as  above  (Fig.  41);  scopa  pale  or  dark  .14 
14a.  Scopa  entirely  pale;  abdomen  largely  reddish  . ...  15 
b.  Scopa  entirely  black  or  pale  on  tibia  and  brownish  on 

basitarsus;  abdomen  dark  16 

15a.  Mesepistemal  pubescence  dark  brownish;  disc  of  sec- 
ond tergite  with  distinct,  minute,  close  punctures;  apex 

of  pygidial  plate  narrowly  truncate  

gelida,  new  species  (part) 

b.  Mesepistemal  pubescence  pale,  at  least  in  part;  disc  of 
second  tergite  without  obvious  minute  punctures,  but 
with  a few  scattered,  moderate,  shallow  punctures;  apex 

of  pygidial  plate  acute  fusciventris  Mocsary 

1 6a.  Pubescence  of  first  and  second  terga  and  of  scopa  black- 
ish; clypeal  punctures,  except  along  impunctate  mid- 
line, uniformly  subcontiguous  on  disc  

obsoleta  (Lepeletier) 

b.  Pubescence  of  adbominal  terga  pale  (brownish  golden 
on  second  segment);  tibial  scopa  mostly  pale;  punctures 

of  clypeal  disc  very  irregularly  spaced 

gelida,  new  species  (part) 

Centris  ( Me/anocentris ) agilis  F.  Smith 

Figure  43 

Centris  agilis  F.  Smith,  1874:361.  3. 

Centris  ignita  F.  Smith,  1874:362.  2.  NEW  SYNONYMY. 
Centris  bakeri  Friese,  1912:199.  <3.  Preoccupied. 

Centris  bakerel/a  Friese,  1913:89.  New  name  for  C.  bakeri 
Friese,  1912,  not  C.  bakeri  Cockerell,  1912.  NEW  SYN- 
ONYMY. 

Epicharis  cisnerosi  Cockerell,  1949:180.  2.  NEW  SYNON- 
YMY. 

F.  Smith  (1874)  described  the  two  sexes  as  C.  agilis  and  C. 
ignita.  This  is  a common  species  in  Mexico  and  one  that  is 
variable  in  the  color  of  the  abdominal  integument.  In  both 
sexes,  the  abdominal  terga,  beyond  the  blackish  first  segment, 
may  be  wholly  ferruginous.  In  these  specimens  the  dense, 
short,  simple  discal  hairs  appear  to  be  yellowish  red.  In  other 
specimens,  one  or  more  of  the  following  three  segments  may 
be  blackish  and  when  this  is  the  case,  the  discal  hairs  appear 
to  be  yellowish  or  somewhat  whitish,  imparting  a distinctly 
“frosted”  appearance. 

Friese’s  C.  bakeri  and  Cockerell’s  Epicharis  cisnerosi  are 
based  on  specimens  with  dark  tergites.  Although  I have  seen 
no  type  material  of  C.  bakeri,  the  description  matches  well 
the  characteristics  of  dark  males  of  C.  agilis.  I have  examined 
the  type  of  Epicharis  cisnerosi ; it  is  a normal,  dark  female 
of  C.  agilis. 

NEW  RECORDS 

MEXICO,  CHIAPAS:  12,  10  mi.  NW  Comitan,  9 Aug.  1963 
(F.D.  Parker  and  L.A.  Stange;  UCD);  12,  Municipio  Ocozo- 
cautla,  El  Aguacero  de  Derna,  762  m elev.,  1 Sept.  1976 
(D.E.  and  J.A.  Breedlove;  CAS);  12,  1833,  Municipio  Angel 
Albino  Corzo,  Rio  Custepec,  below  Finca  Gadow,  853  m 
elev.,  12  Sept.  1976  (D.E.  and  J.A.  Breedlove;  CAS);  1 <3, 
Sumidero,  Tuxtla  Gutierrez,  17  Aug.  1964  (E.  Fisher  and  D. 


Verity;  LACM)  13,  82  mi.  W Tuxtla  Gutierrez,  2100  ft.  elev., 
30  Aug.  1957  (H.A.  Scullen;  ORSU);  222,  NW  slope  Cerro 
Baul,  1768  m elev.,  W of  Rizo  de  Oro,  12  Oct.  1979  (D.E. 
and  J.A.  Breedlove;  CAS).  COLIMA:  13,  10  mi.  W Colima, 
1 Aug.  1954  (M.  Cazier,  W.  Gertsch,  Bradts;  AMNH). 
GUERRERO:  13,  4 mi.  W Chilpancingo,  1530  m elev.,  27 
Aug.  1977  (E.I.  Schlinger;  UCB).  JALISCO:  12,  25  mi  W 
Guadalajara,  4700  ft.  elev.,  29  Sept.  1957  (H.A.  Scullen; 
ORSU);  222,  5.6  km  E Plan  de  Barranca,  914  nr  elev.,  25 
Sept.  1976  (C.D.  George  and  R.R.  Snelling;  LACM),  on  Cas- 
sia sp.;  12,  3.4  km  E Plan  de  Barranca,  960  m elev.,  25  Sept. 
1976  (C.D.  George  and  R.R.  Snelling;  LACM),  on  Cosmos 
sulphureus.  MICHOACAN:  222,  Tuxpan,  6550  ft.  elev.,  19 
Sept.  1957  (H.A.  Scullen;  ORSU).  MORELOS:  13,  no  lo- 
cality or  date  (Crawford;  LACM);  13,  7 mi.  NE  Yautepec, 
4000  ft.  elev.,  18  Aug.  1962  (Univ.  Kans.  Mex.  Exped.; 
UKAN)  on  Leguminosae;  12,  0.6  km  S Teacalco,  1021  m 
elev.,  1 6 Sept.  1976  (C.D.  George  and  R.R.  Snelling;  LACM), 
on  Crota/aria  incana;  13,  Lake  Tequesquitengo,  5000  ft.  elev., 
13  Sept.  1957  (H.A.  Scullen;  ORSU).  NAYARIT:  222,  8 mi. 
N Tepic,  1 Sept.  1962  (D.H.  Janzen;  LACM).  OAXACA:  2 33, 
El  Camaron,  20  Aug.  1959  (L.A.  Stange  and  A.S.  Menke; 
UCB);  233,  48  mi.  E La  Ventosa,  21  July  1963  (J.  Doyen; 
UCB);  12,  near  Pinotepa  Nacional,  200  m elev.,  12  Oct.  1975 
(J.L.  Neff;  LACM),  on  Legume  7 1 23;  222,  4 mi.  W Zanatepec, 
7 Oct.  1975  (J.L.  Neff;  LACM),  on  Cassia  7070;  12,  Mixtla, 
5600  ft.  elev.,  22  Aug.  1963  (Scullen  and  Bolinger;  ORSU). 
PUEBLA:  12,  16.1  km  NW  Izucar  de  Matamoros,  1250  m 
elev.,  1 7 Sept.  1 976  (C.D.  George  and  R.R.  Snelling;  LACM), 
on  Caesalpinia  cacalaco:  322,  22  km  NW  Izucar  de  Mata- 
moros, 1 158  m elev.,  21  Sept.  1976  (C.D.  George  and  R.R. 
Snelling;  LACM),  on  Cassia  laevigata.  SINALOA:  622,  14 
mi.  SE  Espinal,  14  Sept.  1964  (A.E.  Michelbacher;  UCB); 
222,  33  km  SE  Esquinapa,  1 1 Sept.  1 974  (E.M.  Fisher;  LACM). 
VER.4  CRUZ:  13,  Catemaco,  6 Oct.  1976  (E.  Barrera;  LACM). 
HONDURAS:  12,  Zamorano,  2 Nov.  1946  (G.  Cisneros; 
USNM)  (type  of  Epicharis  cisnerosi).  COSTA  RICA,  SAN 
JOSE.  12,  San  Jose,  no  date  (M.  Valerio,  USNM). 

Centris  ( Melanocentris ) agiloides,  new  species 

Figures  20-23 

DIAGNOSIS 

Abdomen  ferruginous  in  both  sexes,  second  tergite  dull,  sub- 
contiguously  punctate;  male  with  clypeus  narrow,  pubescence 
of  thoracic  dorsum  blackish  except  on  metanotum  and  meta- 
femur without  ventral  ridge;  female  with  narrowly  truncate 
pygidial  plate  and  pale  scopa. 

DESCRIPTION 

HOLOTYPE  MALE.  Measurements  (mm).  Head  width 
6.41  (6.00-6.62);  head  length  4.97  (4.56-5.03);  wing  length 
15.0  (14.0-16.5);  total  length  19.0  (17.0-21.0). 

Head.  1.29  (1.29-1.37)  times  broader  than  long;  occipital 
margin  slightly  concave  between  tops  of  eyes;  ocelli  well 
anterior  to  occipital  margin;  inner  orbits  moderately  con- 
vergent above,  upper  frontal  width  0.65  (0.62-0.71)  times 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  27 


Figures  20-23.  Centris  (Melanocentris)  agiloides,  male  seventh  and  eighth  sternites  and  genitalia  (ventral  and  dorsal  views).  Scale  line  = 1.00 
mm. 


lower  frontal  width.  Mandible  stout,  tridentate,  inner  tooth 
acute  and  nearly  as  large  as  second  tooth.  Labrum  about  1.5 
times  wider  than  long,  apex  subacute;  disc  shiny  between 
dense  to  subcontiguous  moderate  punctures.  Clypeus  narrow, 
1.1-1. 2 times  wider  than  long;  median  impunctate  line  nar- 
row and  poorly  defined,  basal  area  of  disc  moderately  shiny, 
roughened  and  with  sparse  fine  punctures,  distal  one-half, 


more  or  less,  shiny  and  weakly  or  not  roughened  and  with 
close  to  dense,  fine  punctures.  Frons  and  preoccipital  area 
moderately  shiny  between  dense  to  subcontiguous  fine  punc- 
tures, except  usual  nearly  impunctate  areas  adjacent  to  ocelli; 
gena  shiny  between  sparse  to  close  fine,  punctures.  Interan- 
tennal  distance  1.62  (1.50-1.77)  times  antennal  socket  di- 
ameter; antennocular  distance  0.46  (0.32-0.47)  times  anten- 


28  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


nal  socket  diameter;  scape  stout,  2.05  ( 1 .90-2.2 1 ) times  longer 
than  wide;  scape  length  0.75  (0.69-0.76)  times  length  of  first 
flagellar  segment;  first  flagellar  segment  longer  than  following 
three  segments  combined,  5.94  (5.00-5.53)  times  length  of 
second.  Interocellar  distance  1.35  (1.26- 1.47)  times  diameter 
of  anterior  ocellus;  ocellocular  distance  0.38  (0.30-0.50)  times 
diameter  of  anterior  ocellus;  ocelloccipital  distance  2.22  (2.16- 
2.50)  times  diameter  of  anterior  ocellus. 

Thorax.  Mesoscutum  moderately  shiny  between  dense  to 
subcontiguous,  fine  punctures  which  are  a little  more  sepa- 
rated posteromedially;  scutellum  with  a low,  rounded  em- 
inence on  either  side,  moderately  shiny  between  dense  to 
subcontiguous,  fine  punctures,  shiny  along  midline;  meta- 
notum  dull,  conspicuously  tessellate  and  with  scattered  fine 
punctures;  mesepisternum  and  metepistemum  moderately 
shiny,  punctures  subcontiguous  to  contiguous,  fine,  and  shal- 
low. Basal  area  of  propodeum  moderately  shiny,  tessellate 
between  sparse,  fine  punctures;  side  and  posterior  face  sim- 
ilar, but  punctures  dense  to  subcontiguous.  Metafemur  stout, 
about  1.9  times  longer  than  thick;  thickest  at  basal  one-third; 
without  ventral  ridge;  metabasitarsus  weakly  curved,  about 
four  times  longer  than  wide. 

Abdomen.  Dorsum  of  first  tergite  moderately  shiny,  weakly 
tessellate  between  sparse,  fine  punctures;  remaining  tergites 
dull  and  sharply  tessellate  between  subcontiguous,  moderate 
punctures,  sparser  on  fourth  and  following  segments;  seventh 
tergum  weakly  bilobate  at  apex. 

Terminalia.  Distal  process  of  seventh  sternite  (Fig.  20) 
short,  not  well  differentiated  from  disc;  apical  emargination 
broad  and  shallow;  hairs  sparse,  mostly  short  and  weakly 
plumose  or  simple.  Median  expansion  of  eighth  sternite  (Fig. 
21)  evenly  rounded;  apex  slightly  convex;  hairs  abundant, 
long,  conspicuously  plumose.  Dorsal  process  of  gonocoxite 
(Figs.  22,  23)  nearly  triangular;  gonostylus  nearly  reaching 
level  of  apex  of  penis  valve;  ventral  lobe  of  penis  valve  only 
slightly  extended  laterad  of  dorsal  lobe. 

Pilosity.  Blackish  brown  on  front  and  top  of  head,  on  tho- 
racic dorsum,  upper  mesepisternum,  anterior  and  middle 
legs,  and  first  tergite;  hairs  on  discs  of  second  and  third  ter- 
gites very  short,  simple,  decumbent,  longer  and  more  erect 
on  following  segments,  with  some  very  long,  plumose,  red- 
dish brown  hairs,  especially  laterad;  hairs  on  underside  of 
head,  posterior  margin  of  scutellum,  metanotum,  and  re- 
mainder of  thorax  light  brown.  Metafemur  and  metabasi- 
tarsus with  scopa-like  yellowish  hairs. 

Color.  Head  and  thorax  blackish  brown,  legs  and  antenna 
more  rufescent.  Large  labral  mark,  large  discal  spot  on  clyp- 
eus  and  narrow  paraocular  mark  all  very  pale  yellowish.  Ab- 
domen dull  ferruginous.  Wings  dark  brown,  veins  and  stigma 
blackish. 

FEMALE.  Measurements  (mm).  Head  width  6.56-7.08; 
head  length  4.82-5.13;  wing  length  14.5-16.0;  total  length 
20.0-24.5 

Head.  1.26-1.38  times  broader  than  long;  ocelli  well  an- 
terior to  nearly  flat  occipital  margin  in  frontal  view;  inner 
orbits  moderately  convergent  above,  upper  frontal  width  0.84- 
0.90  times  lower  frontal  width.  Mandible  quadridentate,  in- 
ner tooth  acute  and  slightly  larger  than  subbasal  tooth.  La- 


brum  about  twice  broader  than  long,  apex  broadly  rounded; 
disc  shiny  between  subcontiguous  to  contiguous,  moderate 
punctures.  Clypeus  mostly  moderately  shiny,  but  disc  with 
variable  area  which  is  conspicuously  shiny  (up  to  distal  two- 
thirds  of  median  area);  median  impunctate  line  poorly  de- 
fined; disc  with  sparse  moderate  punctures  and  a few  low, 
irregular  rugulae  which  are  usually  convergent  distad  or  curved 
mesad  at  their  lower  ends.  Punctation  of  frons  and  occipital 
area  as  described  for  male,  but  interspaces  (including  ocellar 
area)  moderately  shiny  and  conspicuously  tessellate;  gena  as 
described  for  male.  Interantennal  distance  2.00-2.31  times 
antennal  socket  diameter;  scape  robust,  1 .98-2. 1 3 times  longer 
than  wide;  scape  length  0.71-0.77  times  length  of  first  fla- 
gellar segment;  first  flagellar  segment  slightly  longer  than  fol- 
lowing three  segments  combined,  4.52-5.40  times  length  of 
second.  Interocellar  distance  1.55-1.72  times  diameter  of 
anterior  ocellus;  ocellocular  distance  1.08-1.21  times  di- 
ameter of  anterior  ocellus;  ocelloccipital  distance  1.71-2.00 
times  diameter  of  anterior  ocellus. 

Thorax.  As  described  for  male,  but  scutellum  tessellate 
and  slightly  shiny,  except  in  basal  middle  where  it  is  shiny. 
Basitibial  plate  slender,  apex  narrowly  rounded. 

Abdomen.  Dorsal  face  of  first  tergite  moderately  shiny  and 
weakly  tessellate  between  close  to  dense,  moderate  punctures; 
second  and  following  terga  dull,  contiguously  and  finely  punc- 
tate to  tergal  margins;  apex  of  pygidial  plate  narrowly  trun- 
cate. 

Pilosity.  On  head  and  thorax,  as  described  for  male,  but 
hairs  of  metanotum  and  propodeum  often  very  pale  brown- 
ish; scopa  yellowish  white.  Hairs  on  dorsal  face  of  first  tergite 
erect,  dark,  plumose;  dark,  very  short,  appressed,  and  simple 
on  disc  of  second  segment,  becoming  progressively  longer 
and  less  appressed  on  following  segments;  fourth  and  fifth 
terga  with  a few  suberect  to  erect  dark  bristles  on  either  side; 
prepygidial  fimbria  dark  reddish  brown. 

Color.  As  in  male,  but  without  pale  face  marks. 

TYPE  MATERIAL 

Holotype  male:  Finca  La  Selva,  500  ft.  elev.,  near  Puerto 
Viejo,  Heredia  Prov.,  COSTA  RICA,  3 June  1975  (D.R. 
Perry),  on  Dipteryx  panamensis,  0703-1 100,  in  Natural  His- 
tory Museum  of  Los  Angeles  County.  Allotype:  same  locality 
and  collector,  6 May  1979,  on  Dussia  sp.,  1045-1 1 1 5 (LACM). 
Paratypes  (all  COSTA  RICA):  1 1 <33,  69$,  same  data  as  allo- 
type (LACM);  13,  same  locality  and  collector,  4 June  1975, 
on  Dipteryx  panamensis,  1 100-1400  (LACM);  13,  same  lo- 
cality and  collector,  14  June  1975,  on  Dipteryx  panamensis, 
0830-1030  (LACM);  13,  same  locality  and  collector,  24  July 
1978  (LACM),  on  Hymenolobium  sp.;  13,  8 km  S Puerto 
Viejo,  Heredia  Prov.,  28-29  May  1971  (P.A.  Opler;  UCB), 
on  Ipomaea  sp.,  pink  fl.;  13,  Zapote  de  Upala  (vie.  Bijagua), 
Alajuela  Prov.,  19  May  1972  (F.  Cordero;  UCB);  1$,  Dulce 
Nombre,  Cartago  Prov.,  25  Aug.  1967  (R.W.  McDiarmid; 
LACM). 

ADDITIONAL  SPECIMENS  (not  paratypes) 

MEXICO,  CHIAPAS:  13,  Mahosik’,  Tenejapa,  4800  ft.  elev., 
9-12  July  1966  (D.E.  Breedlove  and  J.  Emmel;  CAS).  SAN 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  29 


LUIS POTOSI:  13,  26  mi.  SW  Tamazunchale,  4-5  July  1964 
(E.  Fisher  and  D.  Verity;  LACM);  12,  15  mi.  Xililla,  1350 
m elev.,  15  Aug  1977  (E.I.  Schlinger;  UCB).  VER.4  CRUZ: 
12,  Pehuela,  13  Sept.  1974  (M.  Sousa;  LACM);  422,  same 
locality,  2 Aug.  1974  (A.  Delgado  S.;  LACM),  on  “C.  Doylei 
0800-1002;  12,  “Rig.  de  Cordoba  (A.  Genin;  MNHN).  COS- 
TA RICA,  SAN  JOSE:  2SS,  San  Jose,  no  date  (M.  Valerio; 
USNM).  PANAMA,  PANAMA:  13,  Cerro  Azul,  N of  To- 
cumen,  28  Apr.  1958  (W.J.  Hanson;  UKAN);  13,  Curundu, 
19  May  1981  (R.W.  Brooks;  RWB),  on  Genipa  americana. 

ETYMOLOGY 

Combines  the  Latin  suffix  -oides  (resembling)  to  agilis,  be- 
cause of  the  superficial  resemblance  to  C.  agilis. 

DISCUSSION 

Variation  in  non-meristic  characters  is  negligible.  In  some 
specimens,  particularly  those  of  Mexico,  the  pubescence  of 
the  metanotum  and  propodeum  is  not  conspicuously  paler 
than  that  of  the  thoracic  dorsum.  In  some  males,  the  posterior 
femur,  tibia,  and  basitarsus  may  be  extensively  reddish,  but 
mostly  dark  in  other  specimens.  The  supraclypeal  mark  may 
be  present  or  absent,  and  when  it  is  present,  it  is  transverse 
and  very  narrow.  Although  the  scape  is  usually  black  in  the 
males  available,  there  is  a distinct  ventral  maculation  in  the 
males  from  Mexico  and  Panama.  One  of  the  Mexican  males 
has  the  clypeus  largely  pale. 

In  the  females  there  is  considerable  variation  in  the  extent 
of  the  shiny  area  on  the  clypeal  disc.  In  most  specimens  the 
shiny  portion  of  the  disc  occupies  about  one-half  the  length 
of  the  disc;  in  a few  this  is  exceeded  (up  to  nearly  the  entire 
length)  and  in  others  reduced  to  the  apicomedian  one-fourth 
of  the  segment.  Similarly,  the  development  of  the  oblique 
rugulae  of  the  clypeal  disc  is  variable,  and  the  rugulae,  while 
never  becoming  a dominant  feature,  are  always  present. 

The  uniformly  subcontiguously  punctate,  dull,  red  abdo- 
men is  diagnostic  for  C.  agiloides.  The  only  species  with  a 
similarly  dull  abdomen  is  C.  sericea  but  in  both  sexes  of  that 
species  the  abdomen  is  black,  the  posterior  legs  are  black- 
haired, the  abdominal  punctures  become  much  finer  toward 
the  margins  of  the  segments,  and  the  punctures  of  the  third 
segment  are  much  finer  than  those  of  the  second. 

Centris  {Melanocentris)  flavilabris  Mocsary 

Centris  flavilabris  Mocsary,  1899:253.  2. 

Centris  flavilabris  var.  boliviensis  Mocsary,  1899:  253.  2. 

This  is  a primarily  South  American  species,  not  previously 
recorded  from  Central  America. 

NEW  RECORDS 

COSTA  RICA,  GUANACASTE:  13,  Volcan  Miravalles,  2 
km  W Rio  Navinjo,  15  Mar.  1973  (P.A.  Opler;  UCB).  PUN- 
TARENAS:  12.  Rincon,  25  Apr.  1975  (C.L.  Hogue;  LACM). 


Centris  ( Melanocentris ) fusciventris  Mocsary 

Figure  15 

Centris  fusciventris  Mocsary,  1899:252.  2. 

Centris  fusciventris  var.  scutellata  Friese,  1 900b:273. 3 2.  NEW 
SYNONYMY. 

Centris  (Melanocentris)  fusciventris:  Moure,  1950:388  (tax.). 
Centris  ( Melanocentris ) fusciventris  scutellata:  Michener, 

1 954: 144  (distr.). 

The  var.  C.  f scutellata  was  described  from  both  sexes  from 
Chiriqui,  Panama.  The  nominate  form  occurs  widely  in  South 
America  (Brazil,  Colombia,  Bolivia,  Venezuela).  Differences 
between  the  two  forms  are  slight  and  fall  within  the  range  of 
variation  seen  in  South  American  material. 

NEW  RECORDS 

COSTA  RICA,  HEREDIA:  322,  13,  Finca  La  Selva,  near 
Puerto  Viejo,  6 May  1979  (D.R.  Perry;  LACM),  on  Dussia 
sp.  PANAMA,  COLON:  233,  Puerto  Pilon,  22  km  NE  Santa 
Rita,  23  May  1982  (D.  Roubik;  ROUB).  DARIEN:  3 33,  Bay- 
ano  Bridge,  184  km  SE  Canglon,  16  May  1980  (D.  Roubik; 
ROUB).  PANAMA:  2 33,  Panama,  Curundu,  17  May  1981 
(R.W.  Brooks,  RWB),  on  Genipa  americana. 

Centris  ( Melanocentris ) gelida,  new  species 

DIAGNOSIS 

Female  only:  scopa  yellowish  on  metatibia,  brown  on  meta- 
basitarsus;  abdomen  mostly  ferruginous,  dorsum  shiny  (ob- 
scured by  pubescence)  between  minute  punctures;  dorsal  face 
of  first  tergite  pubescent  and  minutely  punctate  in  middle; 
pubescence  of  first  and  second  tergites  pale.  Male  unknown. 

DESCRIPTION 

HOLOTYPE  FEMALE.  Measurements  (mm).  Head  width 
7.49  (7.38-7.64);  head  length  5.13  (5.03-5.13);  wing  length 
18.0  (17.5-19.0);  total  length  22.0  (22.0-25.0). 

Head.  1.46  (1.46-1.49)  times  broader  than  long;  occipital 
margin  nearly  flat  in  frontal  view  and  slightly  below  tops  of 
eyes;  ocelli  anterior  to  occipital  margin;  inner  orbits  strongly 
convergent  above,  upper  frontal  width  0.85  (0.8 1-0.84)  times 
lower  frontal  width.  Mandible  stout,  quadndentate.  Labrum 
about  twice  broader  than  long.  Disc  of  clypeus  slightly  de- 
pressed, moderately  shiny  (duller,  obviously  tessellate  latero- 
basad),  with  broad,  poorly  defined  median  impunctate  line, 
punctures  fine  and  irregularly  spaced  a puncture  diameter  or 
more  apart.  Frons  roughened  and  slightly  shiny  between  fine, 
dense  punctures;  vertex  slightly  shiny  and  with  punctures 
minute  and  subcontiguous  in  ocellocular  area,  shiny  between 
dense  to  sparse  fine  punctures  in  preoccipital  area;  gena  shiny 
between  dense  to  close,  minute  punctures.  Interantennal  dis- 
tance 2.04  (2.00-2.24)  times  antennal  socket  diameter;  an- 
tennocular  distance  subequal  to  antennal  socket  diameter; 
scape  stout,  about  twice  longer  than  wide,  scape  length  0.72 
(0.63-0.72)  times  length  of  first  flagellar  segment;  first  fia- 


30  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


gellar  segment  slightly  longer  than  following  three  combined, 
4.33  (4.35-4.65)  times  longer  than  second.  Interocellar  dis- 
tance 1.60  (1.42-1.59)  times  diameter  of  anterior  ocellus; 
ocellocular  distance  1.07  (1.00-1.07)  times  diameter  of  an- 
terior ocellus;  ocelloccipital  distance  1.72  (1.68-1.82)  times 
diameter  of  anterior  ocellus. 

Thorax.  Mesoscutum  and  dorsum  of  scutellum  shiny  be- 
tween fine,  dense  to  close  punctures,  posterior  face  of  scu- 
tellum with  subcontiguous,  fine  punctures;  scutellum  weakly 
depressed  along  ntidline;  metanotum  slightly  shiny,  sharply 
tessellate  between  sparse,  minute  punctures;  mesepisternum 
moderately  shiny  between  subcontiguous  to  dense,  fine  punc- 
tures; basal  area  of  propodeum  slightly  depressed  on  either 
side,  moderately  shiny  and  tessellate  between  sparse,  fine 
punctures;  disc  shinier,  closely  and  more  finely  punctate;  side 
shiny  between  fine,  close  punctures.  Basitibial  plate  about 
twice  longer  than  wide,  secondary  plate  narrowly  rounded  at 
apex,  primary  plate  more  broadly  rounded.  Scopal  hairs  plu- 
mose nearly  to  tips. 

Abdomen.  Summit  of  first  tergite  shiny  between  sparse 
micropunctures  in  middle  and  close  to  sparse,  minute  punc- 
tures laterad;  discs  of  second  and  third  tergites  shiny  between 
dense  to  close,  minute  punctures;  fourth  tergite  less  shiny, 
densely,  finely  punctate;  fifth  tergite  slightly  shiny,  subcon- 
tiguously,  finely  punctate.  Pygidium  narrowly  truncate  at  apex, 
sides  straight,  secondary  plate  acute  at  apex. 

Color.  Color  generally  blackish  brown,  distal  half  of  sec- 
ond, all  of  following  tergites  and  all  sternites  ferruginous; 
tarsi  and  underside  of  flagellum  dark  ferruginous.  Wings 
strongly  brownish,  veins  and  stigma  blackish  brown. 

Pilosity.  Dark  brown  to  blackish  on  head,  thorax,  and  legs, 
except  most  of  metatibial  scopa  yellowish  and  metabasitarsal 
scopa  light  brown;  hairs  of  mesoscutum  very  dense,  con- 
cealing surface;  hairs  of  first  tergite  moderately  long,  erect, 
plumose,  dirty  white;  hairs  of  second  to  fifth  tergites  short, 
simple,  subappressed,  whitish  to  yellowish  brown,  especially 
on  discs  of  second  and  third  segments;  prepygidial  and  py- 
gidial  fimbriae  ferruginous;  sternites  with  sparse  erect  whitish 
hairs,  longer  along  midline  and  forming  weak  distal  fimbriae, 
that  of  fifth  segment  ferruginous. 

TYPE  MATERIAL 

Holotype  female:  Santa  Rita,  10  mi.  E Nahuala,  7100  ft. 
elev..  Dept.  Solola,  GUATEMALA,  3 Sept.  1965  (S.J.  Ar- 
nold), on  Canavalia  villosa,  in  California  Academy  of  Sci- 
ences. Paratypes:  22$,  same  data  as  holotype  (UCB);  12,  25 
mi.  S El  Bosque,  5500  ft.  elev.,  Chiapas,  MEXICO,  5 Sept. 
1965  (S.J.  Arnold;  UCB);  one  paratype  in  LACM,  two  in 
UCB. 

ETYMOLOGY 

The  specific  name  is  a Latin  word  for  "frosted”  and  refers 
to  the  characteristic  appearance  of  the  abdominal  dorsum. 

DISCUSSION 

An  additional  female,  not  a paratype,  is  from  Municipio 
Motozintla,  ridge  between  Cerro  Boqueron  and  Niguivil, 


2438-2743  m elev.,  Chiapas,  MEXICO,  1 5 Dec.  1 976  (D.E. 
and  J.A.  Breedlove;  CAS).  This  specimen  is  assumed  to  be 
conspecific  with  the  type  series,  agreeing  closely  in  most  fea- 
tures. It  does  differ,  however,  in  having  the  abdomen  uni- 
formly blackish,  rather  than  mostly  dull  reddish.  Abdominal 
color  is  somewhat  variable  in  some  members  of  this  sub- 
genus. 

The  only  other  species  in  Central  America  with  a similarly 
“frosted”  abdomen  is  C.  agi/is.  In  C.  agilis  the  disc  of  the 
first  tergite  is  polished  and  nearly  devoid  of  punctures  and 
pubescence  across  most  of  its  breadth  and  the  secondary  plate 
of  the  pygidium  is  extended  distad  as  a cariniform  ridge. 

Centris  ( Melanocentris ) obsoleta  Lepeletier 

Figure  39 

Centris  obsoleta  Lepeletier,  1841:153.  2. 

Centris  melanochlaena  F.  Smith,  1874:360.  3.  NEW  SYN- 
ONYMY. 

Epicharis  zamoranensis  Cockerell,  1949:480.  2.  NEW  SYN- 
ONYMY. 

DISCUSSION 

I have  examined  the  type  of  C.  melanochlaena,  from  Ori- 
zaba, Mexico.  It  corresponds  to  the  concept  of  C.  obsoleta, 
in  the  sense  of  Friese  (1900b)  and  subsequent  workers,  and 
it  is  typical  of  Central  American  males  since  the  clypeus  is 
almost  wholly  yellowish. 

I have  also  seen  the  type  and  one  cotype  of  Epicharis 
zamoranensis.  Cockerell  described  the  thoracic  dorsum  as 
being  devoid  of  hairs;  this  is  true  of  the  type,  but  not  of  the 
cotype.  The  hairs  are  worn  off  in  the  type  specimen.  There 
is  no  doubt  that  E.  zamoranensis  is  conspecific  with  C.  ob- 
soleta, a common  species  in  Central  America  and  northern 
South  America. 

NEW  RECORDS 

MEXICO,  NAYARIT:  12,  1 6 mi.  S Acaponeta,  12  Aug.  1963 
(M.G.  Naumann;  UKAN).  OAXACA:  13,  522,  Salina  Cruz, 
7 Sept.  1965  (D.H.  Janzen;  UKAN);  12,  12  mi.  S Chivela, 
18  Aug.  1959  (A.S.  Menke  and  L.A.  Stange;  UCD);  13,  8 km 
W Tehuantepec,  9-10  Aug.  1974  (E.M.  and  J.L.  Fisher; 
LACM).  PUEBLA:  12,  16.1  km  S Izucarde  Matamoros,  1280 
m elev.,  17  Sept.  1976  (C.D.  George  and  R.R.  Snelling; 
LACM),  on  Caesalpinia  cacalaco.  SINALOA:  12,  35  km  SE 
Esquinapa,  1 1 Sept.  1974  (E.M.  Fisher;  LACM).  HONDU- 
RAS: 222,  Zantorano,  2600  ft.  elev.,  5 Nov.  1946  (G.  Cis- 
neros; USNM,  LACM)  (type  and  cotype  E.  zamoranensis). 
COSTA  RICA,  GUANACASTE:  13,  Hacienda  Comelco,  8 
km  NW  Bagaces,  1 9 Nov.  1 97  1 (P. A.  Opler;  UCB),  on  Stach- 
ytarpheta  frantzii. 

Centris  ( Melanocentris ) sericea  Friese 

Figures  24-28 

Centris  sericea  Friese,  1899:41.  2. 

Dr.  R.W.  Brooks  has  examined  the  type  of  C.  sericea,  a 
female  from  an  unspecified  Mexican  locality,  and  found  it 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  31 


to  be  conspecific  with  those  recorded  below.  I have  rede- 
scribed C.  sericea  to  facilitate  its  comparison  with  similar 
species. 

DIAGNOSIS 

Abdomen  black  and  hind  legs  black  pubescent  in  both  sexes. 
Male  metafemur  swollen,  without  ventral  ridge;  inner  man- 
dibular tooth  acute;  clypeus  about  1.2  times  broader  than 
long;  punctures  of  second  tergum  separated  by  a puncture 
diameter  or  more.  Female  with  punctures  of  disc  of  second 
tergum  coarser  than  those  of  apical  zone;  pygidial  plate  with 
margins  strongly  convergent  apicad,  apex  subacute;  thoracic 
pubescence  wholly  dark. 

DESCRIPTION 

MALE.  Measurements  (mm).  Head  width  5.90-6.3 1 ; head 
length  4.46-4.72;  wing  length  15.0-16.0;  total  length  1 8.5— 
20.0. 

Head.  1 .32-1 .34  times  broader  than  long,  occipital  margin, 
in  frontal  view,  slightly  concave  between  tops  of  eyes,  ocelli 
well  anterior  to  occipital  margin;  inner  orbits  moderately 
convergent  above,  upper  frontal  width  0.68-0.70  times  lower 
frontal  width.  Mandible  robust,  tridentate,  inner  tooth  acute 
and  subequal  to  middle  tooth.  Labrum  about  twice  broader 
than  long;  apical  margin  narrowly  rounded;  disc  shiny  be- 
tween subcontiguous.  moderate  punctures.  Clypeus  narrow, 
about  1 .2  times  broader  than  long;  disc  moderately  shiny  to 
shiny  (apical  area),  minutely  roughened  between  close  to 
sparse,  moderate  punctures,  without  distinct  median  im- 
punctate  area.  Frons  slightly  shiny  between  dense,  fine  punc- 
tures, area  in  front  of  anterior  ocellus  and  laterad  of  posterior 
ocelli  impunctate,  tessellate  and  moderately  shiny;  vertex 
moderately  shiny  between  dense,  fine  punctures;  preocciput 
similar  but  punctures  sparse;  gena  moderately  shiny,  punc- 
tures sparse  to  close,  minute  near  eye  grading  to  fine  toward 
gular  margin.  Interantennal  distance  1.75-1.89  times  anten- 
nal socket  diameter;  antennocular  distance  0.48-0.61  times 
antennal  socket  diameter;  scape  stout,  1 .97-2.08  times  longer 
than  broad;  scape  length  0.69-0.77  times  length  of  first  fla- 
gellar segment;  first  flagellar  segment  longer  than  following 
three  combined,  4.41-5.44  times  length  of  second.  Inter- 
ocellar  distance  1.54-1.67  times  diameter  of  anterior  ocellus; 
ocellocular  distance  0.51-0.69  times  diameter  of  anterior 
ocellus  ocelloccipital  distance  2.03—2 . 1 5 times  diameter  of 
anterior  ocellus. 

Thorax.  Mesoscutum  shiny  between  dense  to  subcontig- 
uous, moderate  punctures,  interspaces  tessellate  and  mod- 
erately shiny  in  posterior  middle;  scutellum  slightly  shiny, 
tessellate  between  dense  to  subcontiguous,  moderate  punc- 
tures, a very  low  convexity  on  either  side  of  middle;  meta- 
notum  moderately  shiny  and  sharply  tessellate  between  sparse, 
weak,  fine  punctures;  mes-  and  metepistema  moderately  shiny 
between  dense  to  subcontiguous,  moderate  punctures.  Basal 
area  of  propodeum  moderately  shiny  and  sharply  tessellate 
between  sparse,  fine  punctures;  side  moderately  chiny  and 
roughened  between  close  to  dense,  fine  punctures.  Metafemur 
robust,  about  1.8  times  longer  than  thick,  without  ventral 


ridge;  metabasitarsus  slender,  about  3.6  times  longer  than 
broad. 

Abdomen.  Elevated  basal  area  of  dorsal  face  of  first  tergite 
much  shorter,  in  middle,  than  apical  zone,  slightly  shiny  and 
tessellate  between  sparse,  fine  punctures;  apical  zone  of  first 
tergite  long,  shiny  between  scattered  fine  punctures;  disc  of 
second  tergum  moderately  shiny  between  dense,  moderate 
punctures  which  are  conspicuously  coarser  and  closer  than 
those  of  apical  zone;  disc  of  third  tergite  moderately  shiny 
and  tessellate  between  close,  fine  punctures  which  are  re- 
placed with  fine  setigerous  papillae  on  apical  zone;  fourth  to 
sixth  tergites  moderately  shiny  and  tessellate  between  close, 
slightly  coarser  setigerous  papillae. 

Terminalia.  Seventh  stemite  (Fig.  25)  with  apical  incision 
deeper  and  more  angular  than  in  C.  agiloides  (Fig.  18)  and 
hairs  more  numerous  and  more  conspicuously  plumose. 
Eighth  stemite  (Fig.  26)  more  angularly  expanded  in  middle 
than  in  C.  agiloides  ( Fig.  19).  Gonostylus(Figs.  27,  28)  ending 
well  short  of  level  of  end  of  penis  valve;  ventral  lobe  of  penis 
valve  extended  well  beyond  margin  of  dorsal  lobe. 

Pilosity.  Blackish  brown  on  head,  thorax,  legs,  and  first 
tergite;  second  and  following  tergites  with  whitish  hairs,  very 
short  and  appressed  on  second  segment,  becoming  progres- 
sively longer  and  more  erect  on  succeeding  segments;  golden- 
brown  hairs  laterad  on  fourth  and  following  terga,  on  pygidial 
plate  and  on  sternites. 

Color.  Integument  blackish  brown  on  most  areas;  antenna, 
tegula,  and  legs  dark  reddish  brown;  apical  abdominal  seg- 
ments light  brown.  The  following  pale  yellow:  large  spot  on 
labrum;  clypeus,  except  small  sublateral  brown  spots  near 
base;  transverse  supraclypeal  mark;  paraocular  area,  not  ex- 
tending above  lower  margin  of  antennal  socket;  preapical 
spot  on  underside  of  scape.  Wings  dark  brown,  veins  and 
stigma  blackish. 

FEMALE.  Measurements  (mm).  Head  width  6.67-7.08; 
head  length  4.92-5.33;  wing  length  16.0-17.0;  total  length 
20.5-23.0. 

Head.  1.31-1.37  times  broader  than  long;  occipital  margin, 
in  frontal  view,  nearly  flat  and,  in  center,  slightly  above  level 
of  tops  of  eyes;  ocelli  well  below  occipital  margin;  inner  orbits 
weakly  convergent  above,  upper  frontal  width  0.85-0.93  times 
lower  frontal  width.  Mandible  stout,  quadridentate,  inner 
tooth  acute  and  subequal  to  adjacent  tooth.  Labrum  about 
twice  broader  than  long,  apex  narrowly  rounded,  shiny  be- 
tween subcontiguous,  moderate  punctures.  Clypeus  as  de- 
scribed for  male,  but  about  1.3  times  broader  than  long  and 
median  impunctate  line  a little  more  evident,  but  still  poorly 
defined.  Punctation  of  frons,  occipital  area,  and  gena  as  in 
male.  Interantennal  distance  2.37-2.53  times  antennal  socket 
diameter;  scape  stout,  2.00-2.27  times  longer  than  broad, 
scape  length  0.61-0.73  times  length  of  first  flagellar  segment; 
first  flagellar  segment  longer  than  following  three  segments 
combined,  4.64-5.41  times  longer  than  second.  Interocellar 
distance  1.66-1.95  times  diameter  of  anterior  ocellus;  ocel- 
locular distance  1 .24-1 .45  times  diameter  of  anterior  ocellus; 
ocelloccipital  distance  1.87-1.97  times  diameter  of  anterior 
ocellus. 

Thorax.  Punctation  and  form  as  described  for  male.  Basi- 


32  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


Figures  24-28.  Centris  ( Melanocentris ) sericea,  female  basitibial  plate,  male  seventh  and  eighth  stermtes  and  genitalia  (ventral  and  dorsal 
views).  Scale  line  (25-28)  = 1.00  mm. 


tibial  plate  (Fig.  24)  elongate,  apices  of  primary  and  second- 
ary plates  narrowly  rounded. 

Abdomen.  First  tergum  as  described  for  male,  but  tessellate 


area  limited  to  lateral  areas  of  dorsal  face;  disc  of  second 
tergum  slightly  shiny  and  tessellate  between  fine,  subcontig- 
uous  punctures  which  are  conspicuously  coarser  than  punc- 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  33 


turesofapical  zone;  disc  of  third  tergite  slightly  shiny  between 
dense,  minute  and  scattered,  fine  punctures;  fourth  tergite 
slightly  shiny,  more  sharply  tessellate  between  dense,  fine 
punctures;  fifth  tergite  duller  between  subcontiguous,  fine 
punctures  and  scattered  setigerous  papillae.  Margins  of  py- 
gidial  plate  strongly  convergent  apicad,  apex  narrowly  trun- 
cate or  subacute;  secondary  plate  short  and  broad. 

Pilosity.  As  described  for  male;  prepygidial  fimbria  golden 
brown. 

Color.  As  described  for  male,  but  pale  facial  marks  absent 
and  fifth  tergite  dull  reddish. 

SPECIMENS  EXAMINED 

MEXICO,  MEXICO:  12,  Canon  de  Lobos,  1200  m elev., 
12-13  July  1969  (M.  Sousa;  LACM).  MORELOS:  222,  Yau- 
tepec,  3 1 July  1963  (F.D.  Parker  and  L. A.  Stange;  UCD);  12, 
4 mi.  SW  Yautepec,  3800  ft.  elev.,  2 July  1961  (C.D.  Mich- 
ener;  UKAN),  on  Cassia  sp.;  13,  4.3  mi.  W Yautepec.  4000 
ft.  elev.,  17  Aug.  1962  (Ordway  and  Marston;  UKAN).  NA- 
YARIT:  233,  Km  190,  1200  nr  elev.,  NW  of  Guadalajara,  6 
Sept.  1975  (J.L.  Neff;  LACM),  on  “legume  7046”;  922,  8 mi. 
N Tepic,  1 Sept.  1962  (D.H.  Janzen;  UCB),  on  Crotalaria 
sp. 

DISCUSSION 

Although  the  male  of  C.  sericea  closely  resembles  that  of  C. 
obsoleta,  the  clypeus  is  broader  in  the  latter  species,  at  least 
1.4  times  broader  than  long.  The  inner  margins  of  the  eyes 
are  more  strongly  convergent  above  in  C.  obsoleta,  so  that 
the  distance  between  the  eyes  at  their  upper  ends  is  less  than 
the  distance  between  the  lateral  margins  of  the  two  posterior 
ocelli.  In  C.  sericea  males  the  transocellar  distance  is  less 
than  the  distance  between  the  eyes  at  their  upper  ends.  These 
same  features  will  readily  distinguish  C.  sericea  from  C.  agi- 
lis,  as  will  the  less  shiny,  uniformly  dark  abdomen  and  the 
lack  of  the  pale  hairs  on  the  hind  legs.  From  males  of  C. 
agiloides,  this  species  may  be  separated  by  the  blackish  ab- 
domen, lack  of  pale  pubescence  on  the  hind  legs,  the  mostly 
pale  clypeus,  and  the  generally  more  uniformly  dark  thoracic 
pubescence,  as  well  as  differences  in  abdominal  punctation. 

Females  of  C.  sericea,  by  virtue  of  their  uniformly  dark 
color  and  narrowly  truncate  pygidial  plate  can  only  be  con- 
fused with  those  of  C.  obsoleta.  They  differ  from  that  species, 
however,  in  that  the  disc  of  the  second  tergite  is  densely 
punctate,  the  punctures  conspicuously  coarser  than  those  of 
the  apical  zone  and  those  on  the  disc  of  the  following  segment. 
Additionally,  the  clypeus  is  less  closely  punctate  and  the 
pygidial  plate  is  more  strongly  narrowed  distad  and  less 
broadly  truncate. 

Centris  (. Melanocentris ) p/umipes  F.  Smith 

Centris  plumipes  F.  Smith,  1854:373.  2 3. 

This  species,  originally  described  from  Santarem,  Brazil,  is 
easily  recognized  by  the  features  cited  in  the  key. 

NEW  RECORDS 

COSTA  RICA,  HEREDIA:  3 33,  Finca  La  Selva,  near  Puerto 
Viejo,  6 May  1979  (D.R.  Perry;  LACM),  on  Dussia  sp.;  533, 


same  locality  and  collector,  24  July  1979  (LACM),  on  Hy- 
menolobium  sp. 

Centris  (Melanocentris)  species  A 

Figure  42 

This  is  known  only  from  four  females.  In  this  species,  pu- 
bescence is  dark,  except  light  brownish  on  the  posterior  part 
of  the  scutellum  and  yellowish  on  the  scopa,  the  apex  of  the 
pygidial  plate  is  angularly  emarginate,  and  the  clypeus  is 
conspicuously  maculate.  Although  the  integument  is  gener- 
ally blackish,  the  abdominal  terga  have  weak  metallic  bluish 
reflections. 

SPECIMENS  EXAMINED 

COSTA  RICA,  HEREDIA:  12,  Finca  La  Selva,  near  Puerto 
Viejo,  6 May  1979  (D.R.  Perry;  LACM),  on  Dussia  sp.  PAN- 
AMA, CANAL  ZONE:  12,  Barro  Colorado  Island.  9 July 
1979  (H.  Wolda;  ROUB);  12,  Frijoles,  19  May  1980  (K.E. 
Steiner;  UCD),  on  Byrsomma  crassifolia\  12,  Pipeline  Road, 
3.7  mi.  NW  Gamboa,  9 May  1980  (K.E.  Steiner;  LACM), 
on  B.  crassifo/ia. 

Subgenus  Trachina  Klug 

Trachina  Klug,  1807:226.  Type-species:  Centris  longimana 
Fabricius,  1804;  monobasic. 

Paremisia  Moure,  1945b:406.  Type-species:  “ Paremisia  li- 
neolata  (Lepeletier,  1841)”  = Bombus  similis  Fabricius, 
1804. 

Trachina  is  a member  of  a small  group  of  subgenera,  the 
other  two  being  Heterocentris  and  Hemisiella,  characterized 
by  the  three-segmented  maxillary  palp  in  both  sexes,  the 
swollen  hind  legs  of  the  male  and  the  metabasitarsus  of  the 
male  with  a sharp  posterior  carina  or  ridge  which  terminates 
in  a raised  tooth  (Fig.  18). 

Michener  (1951)  suggested  that  these  three  subgenera  might 
well  be  united  into  a single  genus.  While  there  are  some 
similarities,  it  seems  to  me  that  Trachina,  at  least,  is  distinct 
from  both  Heterocentris  and  Hemisiella.  The  latter  two  sub- 
genera share  a procoxal  modification  which  appears  to  be 
unique  in  Centris.  There  is  a sharply  depressed  groove  along 
the  inner,  ventral  margin  of  the  procoxa;  such  a groove  is 
not  present  in  Trachina  or  the  other  subgenera.  Females  of 
Heterocentris  and  Hemisiella  have  some  scopal  hairs,  es- 
pecially basad  on  the  metatibia,  simple;  in  Trachina  and 
other  subgenera,  the  scopal  hairs  are  conspicuously  plumose. 
Trachina  females  possess  a distinct  secondary  basitibial  plate; 
in  Heterocentris  and  Hemisiella,  the  basitibial  plate  has  a 
basal  convexity,  but  no  secondary  plate. 

Both  sexes  of  Trachina  differ  from  species  of  Hemisiella 
and  Heterocentris  in  possessing  a longitudinal,  median  yellow 
line  on  the  clypeus.  Sometimes,  this  line  may  not  be  evident 
when  the  black  of  the  clypeus  is  reduced  to  a pair  of  subbasal, 
sublateral  spots.  Clypeal  maculae  in  Heterocentris  and  Hemi- 
siella are  usually  present  as  a pair  of  subapical  spots  or  a 
subapical  band  (females,  some  males)  or  else  the  clypeus  is 
yellow  except  across  the  base  (some  males).  Males  of  Tra- 


34  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


Figures  29-37,  left  basitibial  plate  of  female:  29,  Centris  (C. ) flavifrons ; 30,  C.  (C.)  flavofasciata\  31,  C.  (C.)  aethiocesta ; 32,  C.  (C.)  caixensis ; 
33,  C.  (C.)  meaculpa ; 34,  C.  ( Trachina ) eurypalana:  35,  C.  (T.)  fuscata\  36,  C.  (7’.)  dentala\  37,  C.  ( /'. ) xochipillii. 


china  further  differ  in  that  the  apical  lobe  of  the  seventh 
sternite  is  greatly  reduced,  so  that  the  segment  is  essentially 
a transverse  band. 


KEY  TO  NORTH  AMERICAN  TRACHINA 

la.  Male,  antenna  1 3-segmented,  basitibial  plate  absent 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  35 


Figures  38-43,  pygidial  plate  of  female:  38,  Centris  (C.)  flavifrons\  39,  C.  (C.)  flavofasciata:  40,  C.  ( C .)  aelhiocesta\  41,  C.  (Melanocentris) 
obsolete ;;  42,  C.  (M.)  species  A;  43,  C.  (A/.)  agilis. 


36  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


b.  Female,  antenna  1 2-segmented,  basitibial  plate  present 

10 

2a.  Ocellocular  distance  greater  than  diameter  of  anterior 
ocellus;  in  full  frontal  view,  distance  from  anterior  ocel- 
lus to  clypeal  base  a little  shorter  than  distance  from 

anterior  ocellus  to  occipital  margin  3 

b.  Ocellocular  distance  less  than  diameter  of  anterior  ocel- 
lus; in  full  frontal  view,  distance  from  anterior  ocellus 
to  clypeal  base  much  greater  than  distance  from  an- 
terior ocellus  to  occipital  margin  4 

3a.  Thoracic  pubescence  uniformly  blackish,  except  white 
on  scutellum  and  metanotum;  metabasitarsus  with  sharp 
ridge  along  posterior  margin,  terminating  in  sharp  spi- 

niform  process  beyond  midlength  (Fig.  18)  

labiata  Friese 

b.  Flairs  of  mesoscutum  dark  brownish  distally,  base  and 
branches  pale,  so  that  pubescence  appears  “clouded”; 

metabasitarsus  without  posterior  ridge  (Fig.  19) 

longimana  (Fabricius) 

4a.  Abdomen  largely  or  wholly  ferruginous;  if  brownish  or 
blackish,  hairs  of  mesoscutum  red  or  pale  ochreous  . 

5 

b.  First  three,  and  most  of  fourth,  terga  blackish,  apical 
band  of  fourth  and  all  of  fifth  to  seventh  terga  pale 
reddish;  hairs  of  mesoscutum  black  except  for  band  of 

whitish  hairs  across  anterior  one-fourth  

vidua  Mocsary 

5a.  At  least  first  four  terga  dark  brown  to  blackish  (fourth 


may  be  ferruginous  on  apical  margin  6 

b.  Abdomen  largely  or  entirely  ferruginous  7 


6a.  Terga  5-7  dark,  with  glistening  whitish  hairs;  meso- 
scutal  hairs  pale  to  dark  ferruginous;  in  frontal  view, 

occiput  arcuately  raised  above  tops  of  eyes  

si  mi  lis  (Fabricius) 

b.  Terga  5-7  ferruginous,  with  yellowish  to  ferruginous 
hairs;  mesoscutal  hairs  pale  ochreous;  in  frontal  view, 
occiput  flat  or  slightly  concave  between  tops  of  eyes 

xochipillii,  new  species 

7a.  Procoxa  without  distal  spine;  wings  uniformly  light  yel- 
lowish brown  8 

b.  Procoxa  with  slender  distal  spine  (hidden  in  dense  pu- 
bescence); forewing,  basad  of  basal  vein,  clear  and  col- 
orless and  distinctly  brownish  beyond  basal  vein  .... 

eurypatana,  new  species 

8a.  Hairs  of  mesoscutum  pale  ochreous  or  ferruginous; 
metafemur  usually  stout,  subbasal  ventral  process  usu- 
ally tooth-like  (Fig.  57,  58);  second  and  third  terga  mod- 
erately shiny,  disc  finely  tessellate;  middle  and  hind 
legs  with  conspicuous  brownish  to  blackish  pubescence 

9 

b.  Hairs  of  mesoscutum  (and  of  most  of  thorax)  whitish, 
though  often  slightly  dusky  anteriorly;  metafemur  less 
stout,  subbasal  ventral  process  low  and  not  tooth-like 
(Fig.  56);  second  and  third  terga  shiny,  discs  weakly  or 
not  at  all  tessellate;  pubescence  of  legs  whitish  to  pale 
ferruginous,  ferruginous  on  inner  surface  of  metabasi- 
tarsus   heithausi  Snelling 


9a.  Disc  of  seventh  sternite  triangular,  apex  acute  (Fig.  46); 
apical  swelling  of  eighth  sternite  short  and  broad  (Fig. 

47);  basal  lobes  of  penis  valve  short  and  broad  

dentata  F.  Smith 

b.  Disc  of  seventh  sternite  quadrate,  apical  margin  straight 
or  concave  (Fig.  44);  apical  swelling  of  eighth  sternite 
long  and  narrow  (Fig.  45);  basal  lobes  of  penis  valve 

relatively  slender  fuscata  Lepeletier 

10a.  Hairs  of  mesoscutum  dark  brownish  to  blackish  and 
either  (a)  with  a transverse  band  of  white  hairs  across 
front  of  mesoscutum  or  (b)  hairs  of  scutellum  whitish 

11 

b.  Hairs  of  mesoscutum  whitish,  yellowish,  red  or  with 
dark  apices,  but  without  sharply  contrasting  pale  hairs 

anteriorly  or  on  scutellum  12 

11a.  Mesoscutum  with  a band  of  whitish  hairs  across  an- 
terior margin  and  hairs  of  scutellum  and  metanotum 
dark;  first  four  terga  blackish  (fourth  pale  reddish  api- 

cally),  last  two  segments  pale  reddish  

vidua  Mocsary 

b.  Mesoscutum  without  pale  hairs  anteriorly,  but  whitish 
hairs  on  posterior  margin  of  scutellum  and  all  of  meta- 
notum; terga  wholly  ferruginous  labiata  Friese 


1 2a.  First  three  terga  dark  brownish  to  blackish  13 

b.  First  three  terga  ferruginous  14 


13a.  Terga  4 and  5 dark,  margins  colorless,  with  long,  glis- 
tening whitish  hairs;  scopa  black;  mesoscutal  hairs  usu- 
ally dark  reddish;  tergum  4,  across  middle,  with  fine 

dense  piligerous  punctures similis  (Fabricius) 

b.  Terga  4 and  5 ferruginous,  with  yellowish  hairs,  scopa 
pale  yellowish;  mesoscutal  hairs  ochreous;  tergum  4, 
across  middle,  with  sparse  fine  piligerous  punctures 

xochipillii,  new  species 

14a.  Head  width  less  than  5.75  mm;  in  frontal  view,  occiput 
weakly  convex  between  tops  of  eyes;  ocellocullar  dis- 
tance less  than  1.5  x diameter  of  anterior  ocellus;  hairs 

of  thoracic  dorsum  whitish  to  ferruginous 15 

b.  Head  width  more  than  7.0  mm;  in  frontal  view,  occiput 
arcuately  raised  above  tops  of  eyes;  occellocular  dis- 
tance about  twice  diameter  of  anterior  ocellus;  hairs  of 
thoracic  dorsum  with  blackish  shafts  and  pale  branches 

near  base  longimana  (Fabricius) 

15a.  Thoracic  pubescence  ochreous  to  light  brownish  fer- 
ruginous; scape  distinctly  maculate  beneath;  second  and 
third  terga  finely  and  closely  punctate,  surface  mod- 
erately shiny  16 

b.  Thoracic  pubescence  whitish,  tinged  with  pale  brown- 
ish on  mesoscutum;  scape  obscurely,  if  at  all,  maculate; 
second  and  third  terga  shiny  between  fine  punctures 
mostly  separated  by  twice  a puncture  diameter  or  more 

heithausi  Snelling 

16a.  Wings  light  to  medium  yellowish  brown;  lower  margin 
of  secondary  basitibial  plate  strongly  oblique  (Fig.  36) 
or,  if  somewhat  transverse,  posterior  margin  is  contin- 
uous with  that  of  primary  plate  (Fig.  35)  17 

b.  Wings  blackish  brown;  lower  margin  of  secondary  basi- 
tibial plate  transverse,  anterior  margin  curved  basad 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  37 


well  before  anterior  margin  of  primary  plate  (Fig.  34) 

eurypatana,  new  species 

1 7a.  Basitibial  plate  broad,  lower  margin  of  secondary  plate 
usually  transverse  across  middle  portion,  posterior 
margin  continuous  with  that  of  primary  plate  (Fig.  35) 

fuscata  Lepeletier 

b.  Basitibial  plate  narrow,  lower  margin  of  secondary  plate 
strongly  oblique  throughout,  posterior  margin  extended 
over  that  of  primary  plate  (Fig.  36)  . dentata  F.  Smith 

Centris  ( Trachina ) dentata  F.  Smith 

Figures  36,  46,  47,  58 

Centris  dentata  F.  Smith.  1854:374.  <3. 

Centris  proximo  Friese,  1899:45.  <3  2. 

Michener  (1954)  recorded  C.  dentata  from  Panama.  In  ad- 
dition to  material  from  Panama,  I have  seen  several  females 
from  Mixtepec,  Oaxaca,  Mexico,  28  Feb.  1974  (M.  Sousa; 
LACM).  This  appears  to  be  an  uncommon  species  in  Central 
America,  although  widely  distributed  in  South  America;  the 
types  of  both  C.  dentata  and  C.  proximo  were  from  Brazil. 

Centris  ( Trachina ) eurypatana,  new  species 

Figures  34,  48-51,  57 

DIAGNOSIS 

Male  distinguished  from  all  other  Trachina  by  the  presence 
of  a slender  spine  at  the  apex  of  the  procoxa.  Female  separable 
from  other  Trachina  by  the  following  combination:  pubes- 
cence pale  ochreous,  abdomen  ferruginous,  wings  dark  brown, 
posterior  margin  of  secondary  basitibial  plate  extended  be- 
yond that  of  primary  plate. 

DESCRIPTION 

HOLOTYPE  MATE.  Measurements  (mm).  Head  width 
5.90;  head  length  4.10;  wing  length  13.0;  total  length  15.0. 

Head.  1 .44  times  broader  than  long;  occipital  margin  weakly 
convex  in  frontal  view,  ocelli  well  anterior  to  margin;  inner 
orbits  strongly  convergent  above,  upper  frontal  width  0.68 
times  lower  frontal  width.  Mandible  slender,  tridentate,  inner 
tooth  large  and  its  lower  edge  slightly  sinuate.  Labrum  about 
1 .4  times  broader  than  long,  apical  margin  broadly  rounded; 
disc  shiny  between  subcontiguous,  fine  punctures.  Clypeus 
about  1.4  times  broader  than  long;  disc  shiny  across  apical 
margin,  otherwise  slightly  shiny  and  distinctly  tessellate  be- 
tween dense  to  subcontiguous,  fine  punctures,  impunctate 
median  line  narrow  and  slightly  elevated.  Frons  moderately 
shiny  between  dense  to  subcontiguous,  fine  to  moderate 
punctures,  sparsely  punctate  areas  adjacent  to  ocelli  mod- 
erately shiny  and  distinctly  tessellate;  preoccipital  area  shiny 
between  dense  subcontiguous,  fine  to  minute  punctures;  gena 
shiny  between  close  to  dense,  fine  punctures.  Interantennal 
distance  0.82  times  antennal  socket  diameter,  scape  robust, 
1.97  times  longer  than  wide;  scape  length  0.70  times  length 
of  first  flagellar  segment;  first  flagellar  segment  5.63  times 
longer  than  second,  distinctly  longer  than  combined  lengths 
of  second  to  fourth  segments.  Interocellar  distance  1 .90  times 


diameter  of  anterior  ocellus;  ocellocular  distance  0.55  times 
diameter  of  anterior  ocellus;  ocelloccipital  distance  2.84  times 
diameter  of  anterior  ocellus. 

Thorax.  Mesoscutum  shiny  between  subcontiguous,  fine 
punctures  which  become  well  separated  posteromedially; 
scutellum  shiny  between  scattered,  minute  to  fine  punctures; 
metanotum  dull,  sharply  tessellate  and  with  scattered,  minute 
punctures;  mesepisternum  and  metepisternum  shiny  be- 
tween dense  to  subcontiguous,  fine  to  moderate  punctures. 
Basal  area  of  propodeum  shiny  and  weakly  tessellate  between 
sparse  to  close,  moderate  punctures;  side  and  disc  shiny  be- 
tween close  to  dense,  fine  to  moderate  punctures.  Procoxa 
with  narrow,  distal,  spiniform  process;  metafemur  stout,  about 
twice  longer  than  thick,  ventral  process  stout  (Fig.  57);  pro- 
cess on  anterior  margin  of  metatibia  slender,  spiniform; 
metabasitarsus  about  3.2  times  longer  than  broad,  posterior 
ridge  ending  at  about  midlength. 

Abdomen.  Dorsal  face  of  first  tergite  moderately  shiny  and 
tessellate  between  sparse,  minute  punctures;  disc  of  second 
tergite  shiny  and  weakly  tessellate  between  sparse  to  dense, 
fine  punctures,  punctures  of  apical  zone  minute;  third  tergite 
similar,  but  punctures  dense;  fourth  tergite  similar  to  third, 
but  punctures  variably  spaced  from  sparse  to  subcontiguous; 
fifth  tergite  similar  to  fourth,  but  punctures  sparse;  apex  of 
seventh  tergite  distinctly  bilobed,  hidden  under  dense  hairs. 

Terminalia.  Process  of  seventh  sternite  (Fig.  48)  short, 
broad,  apical  margin  convex;  margin  with  long,  plumose 
hairs,  disc  with  a few  short,  simple  hairs.  Eighth  sternite  (Fig. 
49)  moderately  flared  preapically,  apex  acute;  hairs  fine,  plu- 
mose. Dorsal  process  of  gonocoxite  short,  broad,  apex  round- 
ed; distal  tubercles  of  gonostylus  not  visible  in  dorsal  view 
(Figs.  50,  5 1 ). 

Pilosity.  Generally  pale  ochreous,  slightly  brownish  across 
vertex,  on  thoracic  dorsum,  and  on  legs,  darker  on  meso- 
and  metatibiae  and  tarsi;  some  brown  hairs  on  inner  surface 
of  meso-  and  metatibiae  and  basitarsi.  Second  tergite  with 
long,  suberect,  plumose,  blackish  brown  hairs  across  base, 
discs  of  second  to  fourth  terga  with  sparse,  simple,  dark  hairs 
which  become  progressively  longer  and  more  erect  on  suc- 
ceeding segments;  hairs  on  fifth  to  seventh  terga  long,  sub- 
erect to  erect,  slightly  yellowish  to  ferruginous.  Ventral  ab- 
dominal pubescence  dense,  light  yellowish  brown. 

Color.  Head  and  thorax  blackish  brown,  abdomen  ferru- 
ginous; antenna  and  legs  variably  light  to  dark  reddish  brown. 
The  following  yellow:  mandible,  except  apical  teeth;  labrum; 
clypeus,  except  a pair  of  brown  submedian  spots  near  base; 
transverse  supraclypeal  stripe;  paraocular  area,  upper  end 
constricted  and  terminating  on  eye  margin  at  about  midlevel 
of  antennal  socket;  broad  ventral  stripe  on  scape;  dorsal, 
apical  spot  on  profemur;  basal  spot  on  pro-  and  mesotibia. 
Tegula  clear  yellowish-brown.  Wings  clear  and  very  light 
brown  basad  of  vein  M,  darker  brown  distad;  veins  and 
stigma  blackish  brown. 

FEMALE.  Measurements  (mm).  Head  width  5.87-6.10; 
head  length  4.05-4.27;  wing  length  11.5-12.5;  total  length 
15.0-17.0. 

Head.  1.43-1.44  times  broader  than  long;  in  frontal  view, 
occipital  margin  gently  convex,  ocelli  well  anterior  to  margin; 


38  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


46 


Figures  44-47,  seventh  and  eighth  sternites  of  male:  44-45,  Centris  ( Trachina)  fiuscata:  46-47,  C.  (T.)  dentata.  Scale  line  = 0.50  mm. 


inner  orbits  moderately  convergent  above,  upper  frontal  width 
0.87-0.89  times  lower  frontal  width.  Mandible  tridentate, 
inner  tooth  large,  blunt.  Labrum  about  1.5  times  broader 
than  long,  margin  broadly  rounded;  disc  shiny  between  sub- 
contiguous,  fine  to  moderate  punctures.  Clypeus  about  1 .6 
times  broader  than  long,  otherwise  as  described  for  male. 
Frons  and  preoccipital  area  as  described  for  male;  gena  shiny, 
punctures  close  and  minute  near  eye,  becoming  dense  and 
fine  ventrad.  Interantennal  distance  2.79-3.00  times  anten- 
nal socket  diameter;  antennocular  distance  1.61—1.71  times 
antennal  socket  diameter;  scape  stout,  2.03-2. 1 7 times  longer 
than  wide,  scape  length  0.67-0.72  times  length  of  first  fla- 
gellar segment;  first  flagellar  segment  5.63-5.88  times  longer 


than  second,  longer  than  following  three  segments  combined. 
Interocellar  distance  2. 1 4—2.30  times  diameter  of  anterior 
ocellus;  ocellocular  distance  1.33-1.48  times  diameter  of  an- 
terior ocellus;  ocelloccipital  distance  2.80-3.00  times  di- 
ameter of  anterior  ocellus. 

Thorax.  As  described  for  male.  Procoxa  without  ventral 
spine;  basitibial  plate  (Fig.  34)  narrowly  rounded  at  apex, 
posterior  margin  of  secondary  plate  beyond  that  of  primary 
plate. 

Abdomen.  First  two  terga  as  described  for  male,  third  with 
punctures  of  disc  minute  and  sparse,  becoming  more  minute 
and  scattered  in  apical  zone;  fourth  tergum  less  shiny  than 
third,  punctures  very  irregularly  spaced,  moderate  and  some- 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  39 


what  elongate,  close  to  dense.  Pygidial  plate  with  margins 
strongly  convergent  distad,  apex  narrowly  truncate;  second- 
ary plate  obsolete. 

Pilosity.  About  as  described  for  male,  but  some  hairs  on 
disc  of  fourth  tergite  at  least  weakly  plumose;  scopa  pale 
yellowish;  prepygidial  fimbria  light  golden  brown. 

Color.  As  described  for  male,  except  mandibular  apex  more 
extensively  dark,  dark  clypeal  spots  reaching  base,  supra- 
clypeal  spot  greatly  reduced  or  absent  and  legs  approximately 
concolorous  with  abdomen.  Wings  darker  brownish. 


TYPE  MATERIAL 

Holotype  male:  Estacion  Biologia  Chamela,  Jalisco,  MEX- 
ICO, 17  May  1980  (S.H.  Bullock,  #373),  in  Natural  History 
Museum  of  Los  Angeles  County.  Allotype:  same  locality  and 
collector,  7 Apr.  1982  (#922;  LACM).  Paratypes  (all  MEX- 
ICO): 12,  same  locality  and  collector,  12  May  1980  (#372); 
222,  30  mi.  E Villa  Union,  570  m elev.,  Sinaloa,  10  Mar. 
1980  (J.L.  Neff;  NEFF).  One  paratype  each  in  collections  of 
the  Estacion  Biologia  de  Chamela  and  in  the  personal  col- 
lection of  J.L.  Neff,  the  remaining  specimens  in  the  Natural 
History  Museum  of  Los  Angeles  County. 


ETYMOLOGY 

The  specific  epithet  combines  the  Greek  eurys  (broad)  and 
patanas  (plate),  in  allusion  to  the  broad  secondary  basitibial 
plate  of  the  female. 


DISCUSSION 

The  male  is  easily  distinguished  from  all  previously  described 
species  of  Trachina  by  the  presence  of  a distinct  spine-like 
process  at  the  apex  of  the  procoxa.  It  is  otherwise  very  similar 
to  the  males  of  C.  dentata,  C.  fuscata,  C.  heithausi,  and  C. 
xochipillii.  All  of  these  differ  from  males  of  C.  eurypatana 
in  having  lighter  colored  wings,  particularly  that  of  C.  heit- 
hausi, which  further  differs  in  the  paler  pubescence  of  the 
head  and  thorax.  Males  of  C.  dentata  have  the  ventral  tooth 
of  the  metafemur  higher  and  more  slender  and  the  tooth  on 
the  anterior  margin  of  the  metatibia  is  stouter  than  in  C. 
eurypatana.  The  brown,  rather  than  ferruginous,  first  three 
abdominal  terga  will  differentiate  C.  xochipillii  from  C.  eu- 
rypatana. 

The  female  of  C.  eurypatana  is  best  recognized  from  other 
members  of  this  complex  by  the  broad  secondary  basitibial 
plate,  ochreous  pubescence,  dark  wings,  and  wholly  red  ab- 
domen. In  C.  xochipillii  the  first  three  abdominal  segments 
are  brown;  in  C.  heithausi  the  thoracic  pubescence  is  whitish 
and  the  wings  only  faintly  brown;  in  C.  dentata  the  median 
line  of  the  clypeus  is  shiny  and  the  punctures  on  either  side 
are  moderate  rather  than  fine,  and  the  second  tergite  is  duller, 
more  sharply  tessellate  and  closely  punctate.  Since  the  sec- 


ondary basitibial  plate  of  C.  fuscata  does  not  overhang  the 
first,  this  species  is  easily  separated  from  C.  eurypatana. 

Centris  ( Trachina ) fuscata  Lepeletier 

Figures  34,  44,  45 

Centris  fuscata  Lepeletier,  1841:167.  <3. 

Centris  bimaculata  Lepeletier,  1841:168.  2. 

This  species,  also  described  from  Brazil,  is  much  more  com- 
mon in  Central  America  than  the  similar  C.  dentata.  Females 
of  C.  fuscata,  however,  have  a narrow  secondary  basitibial 
plate  (Fig.  35),  a feature  which  will  separate  this  species  from 
superficially  similar  species.  Males  most  closely  resemble 
those  of  C.  dentata,  but  have  the  ventral  tooth  of  the  meta- 
femur short  and  stout;  in  C.  dentata  this  tooth  is  long,  slender, 
and  somewhat  curved. 

I have  seen  Central  American  material  from  Mexico,  Gua- 
temala, Costa  Rica,  and  Panama.  Michener  (1954)  has  also 
recorded  C.  fuscata  from  Panama  and  Lutz  and  Cockerell 
(1920)  cite  it  from  Guatemala. 

Centris  ( Trachina ) heithausi  Snelling 

Figure  56 

Centris  ( Trachina ) heithausi  Snelling,  1974:20-23.  <3  2. 

This  species  was  described  from  many  specimens  from 
Guanacaste  Province,  Costa  Rica.  I have  seen  numerous  ad- 
ditional specimens  from  the  same  general  area.  One  female, 
however,  was  collected  20  km  SSE  of  Chiquimula,  Depto. 
Chiquimula,  Guatemala,  25  Feb.  1966  (D.P.  Gregory;  UCB). 

Centris  ( Trachina ) labiata  Friese 

Figure  18 

Centris  labiata  Friese,  1904:91.  <3. 

Centris  schwarzi  Cockerell,  1919:192.  2.  NEW  SYNONY- 
MY. 

This  is  evidently  not  a common  species.  Friese’s  male  type 
is  from  San  Carlos,  Costa  Rica.  The  type  of  C.  schwarzi  is 
from  Alta  Vera  Paz,  Guatemala.  I have  seen  the  type  of  C. 
schwarzi,  in  the  USNM,  and  it  is  the  same  as  females  I had 
already  associated  with  C.  labiata  males.  Both  sexes  are  char- 
acterized by  the  dark  thoracic  pubescence  (but  pale  on  the 
scutellum  and  metanotum)  and  the  red  abdomen. 

NEW  RECORDS 

MEXICO,  OAXACA:  2<3<3,  20  mi.  E El  Cameron,  21  July 
1956  (J.W.  MacSwain;  UCB);  2<3<3,  19  mi.  W Tequisistlan, 
29  Aug.  1970  (E.M.  and  J.L.  Fisher;  LACM).  VERA  CRUZ: 
1<5,  El  Palmar,  28  Mar.  1954  (D.H.  Janzen;  LACM).  YU- 
CATAN: 12,  Piste,  29  June  1967  (E.C.  Welling;  LACM). 
BELIZE:  12,  no  further  data  (LACM). 

Centris  ( Trachina ) longimana  Fabricius 

Figure  19 

Centris  longimana  Fabricius,  1804:356.  2 <3. 


40  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


Figures  48-51.  Centris  (Track ina)  eurypatana,  male  seventh  and  eighth  sternites  and  genitalia  (ventral  and  dorsal  views).  Scale  line  = 1.00 
mm. 


Centris  personata  F.  Smith,  1874:362.  <3. 

Michener  (1954)  reported  C.  longimana  from  several  Pan- 
amanian localities.  In  addition  to  specimens  from  Panama, 
I have  seen  material  from  Nicaragua  and  Costa  Rica. 

Centris  ( Trachina ) similis  (Fabricius) 

Bombus  similis  Fabricius,  1 804:35 1 . 9. 

Centris  lineolata  Lepeletier,  1841:158.  9. 

Centris  lineolata  castaneiventris  Mocsary,  In  Friese,  1899: 
288. 

Centris  (Paremisia)  similis:  Moure,  1 960b:  1 30-13 1 . 


This  is  a common  species  in  northern  South  America  (Trin- 
idad, Guyana,  French  Guiana)  south  to  Brazil  and  Peru. 
There  are  no  previous  reports  of  its  presence  in  Central 
America.  I have  seen  the  following  Central  American  spec- 
imens. 

NEW  RECORDS 

COSTA  RICA,  PUNTARENAS:  19,  1.8  mi.  W Rincon,  4 
Mar.  1971  (J.P.  Donahue  and  C.L.  Hogue;  LACM).  SAN 
JOSE:  19.  Pozo  Azul,  Junction  Rios  Parrita  and  Candelaria, 
85  m elev.,  9 Dec.  1961  (A.  Wille;  UKAN).  PANAMA. 
PANAMA:  2i3<3,  15  km  E Chepo,  Llano  Carti  Rd.,  18  Jan. 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  41 


56  57  58 


Figures  52-55.  Centris  (Trachina)  xochipillii,  male  seventh  and  eighth  sternites  and  genitalia  (ventral  and  dorsal  views).  Scale  line  = 1.00  mm. 
Figs.  56-58,  outline  of  ventral  metafemoral  process  of  male:  56,  C.  (T.)  heithausr,  57,  C.  (T.)  eurypatana\  58,  C.  (T.)  dentata. 


42  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


1980  (D.  Roubik,  #6;  ROUB);  292,  15,  15  km  NE  Chepo,  19 
Dec.  1980  (D.  Roubik,  #48;  ROUB). 

Centris  ( Trachina ) vidua  Mocsary 

Centris  vidua  Mocsary,  1899:252.  5. 

This  species  was  described  from  Honduras  (San  Pedro  Sula) 
and  has  been  reported  by  Friese  (1900b)  from  Orizaba,  Vera 
Cruz,  Mexico.  I have  seen  the  following  specimens  of  this 
uncommon  bee. 

NEW  RECORDS 

BELIZE:  1 2,  Belize  (no  further  data;  LACM).  COSTA  RICA, 
CARTAGO:  15,  Turrialba,  18  Oct.  1947  (A.  Svihla;  LACM). 
PANAMA,  BOCAS  DEL  TORO:  12,  Almirante,  Sept.  1963 
(LACM).  COLON:  222,  15,  5 km  SW  Colon,  30  Jan.  1980 
(D.  Roubik,  #12;  ROUB). 

Centris  ( Trachina ) xochipillii,  new  species 

Figures  37,  52-55 

DIAGNOSIS 

At  least  first  three  terga  brown,  apical  segments  ferruginous; 
male  ocellocular  distance  less  than  ocellar  diameter,  occipital 
margin  at  most  weakly  convex  in  frontal  view,  thoracic  pu- 
bescence ochreous;  female  with  ochreous  thoracic  pubes- 
cence, posterior  margin  of  secondary  plate  of  basitibial  plate 
overhanging  that  of  primary  plate,  scopa  yellowish. 

DESCRIPTION 

HOLOTYPE  MALE.  Measurements  (mm).  Head  width 
5.23  (5.13-5.74);  head  length  3.79  (3.59-4.05);  wing  length 
13.5  (12.0-14.0);  total  length  15.0  (13.0-17.0). 

Head.  1 .38  ( 1 .38— 1.51)  times  broader  than  long;  in  frontal 
view,  occipital  margin  flat  or  slightly  convex,  ocelli  well  be- 
low margin;  inner  eye  margins  strongly  convergent  above, 
upper  frontal  width  0.74  (0.69-0.75)  times  lower  frontal  width. 
Mandible  tridentate,  inner  tooth  large,  its  lower  margin  slightly 
concave  in  outline.  Labrum  about  1 .7  times  wider  than  long, 
apical  margin  broadly  rounded;  disc  shiny  between  subcon- 
tiguous,  fine  to  moderate  punctures.  Clypeus  about  1.6  times 
broader  than  long;  disc  dull  and  densely  tessellate  at  base, 
shiny  and  weakly  tessellate  on  about  distal  one-fourth,  me- 
dian line  raised  and  impunctate,  disc  otherwise  densely  to 
subcontiguously  punctate,  punctures  fine  to  moderate.  Frons 
and  ocellar  area  closely  punctate,  except  usual  nearly  im- 
punctate areas  near  ocelli;  postocellar  area  varying  from 
moderately  shiny  between  minute,  dense  punctures  imme- 
diately behind  ocelli,  to  shiny  between  sparse  to  close,  mod- 
erate punctures  at  posterior  margin;  gena  shiny  between  sparse 
to  close  punctures,  minute  adjacent  to  eye  and  grading  to 
fine  over  most  of  area.  Interantennal  distance  2.30  (2.00- 
2.56)  times  antennal  socket  diameter;  antennocular  distance 
0.85  (0.63-0.88)  times  antennal  socket  diameter;  scape  stout, 
1.90  (1.7 6—2.03)  times  longer  than  wide,  scape  length  0.63 
(0.63-0.72)  times  length  of  first  flagellar  segment;  first  fla- 


gellar segment  longer  than  following  three  segments  com- 
bined, 5.63  (5.24-6.13)  times  length  of  second  segment.  In- 
terocellar  distance  2. 1 5 ( 1 .89-2. 1 1 ) times  diameter  of  anterior 
ocellus;  ocellocular  distance  0.70  (0.57-0.78)  times  diameter 
of  anterior  ocellus;  ocelloccipital  distance  2.85  (2.50-2.96) 
times  diameter  of  anterior  ocellus. 

Thorax.  Mesoscutum  shiny  between  dense,  fine  punctures; 
dorsal  face  of  scutellum  shiny,  punctures  sparse  and  minute 
in  center,  becoming  close  and  line  laterad,  dense  and  fine  on 
posterior  face;  metanotum  slightly  shiny,  sharply  tessellate 
between  scattered,  minute  punctures;  mesepisternum  shiny 
between  dense,  fine  punctures;  metespisternum  similar  but 
punctures  more  crowded  toward  posterior  margin  and  lower 
one-third  with  punctures  minute  and  scattered.  Basal  area 
shiny  between  sparse,  fine  punctures  in  middle,  becoming 
dense  laterad;  side  of  propodeum  similar  but  punctures  scat- 
tered to  close.  Procoxa  aspinose;  metafemur  robust,  about 
1.8  times  longer  than  thick,  ventral  process  stout;  anterior 
tooth  of  metatibia  acute,  moderately  stout;  metabasitarsus 
about  3.5  times  longer  than  wide,  posterior  carina  ending  at 
about  midlength  in  slender  tooth. 

Abdomen.  Dorsal  face  of  first  tergum  moderately  shiny  and 
tessellate  between  sparse,  minute  punctures;  disc  of  second 
tergum  shiny  between  sparse,  minute  punctures  which  extend 
across  apical  zone  nearly  to  margin;  disc  of  third  tergum 
similar,  but  punctures  a little  larger  and  impunctate  margin 
broader;  fourth  tergum  shiny  and  weakly  tessellate  between 
sparse,  moderate  punctures;  fifth  and  sixth  terga  similar,  but 
a little  more  distinctly  tessellate. 

Terminalia.  Seventh  sternite  (Fig.  52)  with  distal  process 
longer  and  more  narrowly  rounded  than  in  C.  eurypatana 
(Fig.  48)  and  fewer  long  marginal  hairs.  Apical  swelling  of 
eighth  sternite  (Fig.  51)  shorter  than  in  C.  eurypatana  (Fig. 
49),  apex  less  acute.  Dorsal  process  of  gonocoxite  (Fig.  55) 
narrower  and  more  acute  than  in  C.  eurypatana ; tubercles 
at  apex  of  gonostylus  visible  in  dorsal  view. 

Pilosity.  Pubescence  generally  ochreous,  whitish  on  head, 
especially  on  gena;  brownish  across  vertex,  anteriorly  on  me- 
sobasitarsus,  externally  on  metatibia  and  metabasitarsus; 
reddish  brown  on  remainder  of  hind  legs;  discs  of  second  to 
fourth  terga  with  hairs  simple,  blackish  and  appressed  on 
second,  longer  and  more  erect  on  third,  some  suberect  on 
fourth;  fifth  with  erect  hairs,  some  plumose,  mixed  ferrugi- 
nous and  black;  sixth  and  seventh  terga  with  hairs  mostly 
plumose,  pale  ferruginous,  brownish  laterad. 

Color.  Blackish;  first  three  terga,  and  most  of  fourth,  dark 
reddish  brown;  distal  margin  of  fourth,  and  all  of  following 
terga,  ferruginous;  sternites  reddish;  legs  medium  to  light 
reddish  brown.  The  following  dull  yellow:  most  of  mandible; 
labrum;  narrow  median  stripe  and  transverse  distal  band 
(broader  laterad)  on  clypeus;  paraocular  area,  constricted 
above  and  ending  on  eye  margin  at  about  midlength  of  an- 
tennal socket;  linear  mark  on  underside  of  scape;  small  basal 
spot  on  pro-  and  mesotibiae.  Wings  light  brown,  a little  dark- 
er apicad;  veins  and  stigma  dark  brown. 

FEMALE.  Measurements  (mm).  Head  width  5.79-6.10; 
head  length  4.21-4.36;  wing  length  1 1.0-13.0;  total  length 
13.0-17.0. 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  43 


Head.  1.36-1.42  times  broader  than  long;  in  frontal  view 
occiput  nearly  flat  and  ocelli  well  anterior  to  margin;  eyes 
weakly  convergent  above,  upper  frontal  width  0.88-0.9 1 times 
lower  frontal  width.  Mandible  tridentate,  inner  tooth  large, 
subtruncate.  Labrum  about  1 .6  times  broader  than  long,  apex 
broadly  rounded;  shiny  between  subcontiguous,  moderate 
punctures,  but  with  distinct,  slightly  convex  basal  zone  vir- 
tually free  of  punctures.  Clypeus  about  1.7  times  broader 
than  long,  otherwise  about  as  described  for  male,  but  im- 
punctate  median  line  moderately  shiny  and  moderate-sized 
punctures  clearly  dominant.  Frons,  occipital  area,  and  gena 
about  as  described  for  male.  Interantennal  distance  2.43- 
2.77  times  antennal  socket  diameter,  antennocular  distance 
1 .47-1 .68  times  antennal  socket  diameter;  scape  robust,  scape 
1.83-2.06  times  longer  than  broad;  scape  length  0.70-0.76 
times  length  of  first  flagellar  segment;  first  flagellar  segment 
longer  than  following  three  segments  combined,  5.00-5.41 
times  length  of  second  segment.  Interocellar  distance  2.06- 
2.24  times  diameter  of  anterior  ocellus;  ocellocular  distance 
1.39-1.48  times  diameter  of  anterior  ocellus;  ocelloccipital 
distance  2.50-2.83  times  diameter  of  anterior  ocellus. 

Thorax.  Punctation  as  described  for  male.  Basitibial  plate 
(Fig.  37)  long,  apex  subacute;  posterior  margin  of  secondary 
plate  extending  beyond  that  of  primary  plate. 

Abdomen.  First  tergum  similar  to  that  of  male,  but  with  a 
few  moderate  punctures  across  base  of  dorsal  face;  second 
to  fifth  terga  about  as  described  for  male.  Pygidial  plate 
V-shaped,  apex  narrowly  rounded,  secondary  plate  indis- 
tinct. 

Pilosity.  Generally  as  described  for  male,  but  only  first 
three  terga  dark  reddish  brown,  last  three  ferruginous;  yel- 
lowish marks  of  face  as  in  male,  but  scape  entirely  dark,  or 
with  a small,  obscure,  yellowish  blotch. 

TYPE  MATERIAL  (all  Oaxaca,  MEXICO) 

Holotype  male,  allotype,  233,  792  paratypes;  Tehuantepec. 
18  Feb.  1954  (R.R.  Dreisbach),  in  University  of  Kansas, 
Snow  Entomological  Museum.  Additional  paratypes:  12,  20 
mi.  W Tehuantepec,  18  Feb.  1954  (R.R.  Dreisbach;  UKAN); 
13,  5 mi.  W Tehuantepec,  7 Apr.  1953  (E.I.  Schlinger;  UCB); 
833,  35  mi.  N Tehuantepec,  2600  ft.  elev.,  5 Feb.  1966  (D. 
Bolinger;  ORSU);  13,  20  mi.  E Juchitan  jet.,  500  ft.  elev.,  28 
Jan.  1965  (D.  Bolinger;  ORSU).  Two  male  and  two  female 
paratypes  in  LACM,  remainder  returned  to  their  respective 
collections. 

ETYMOLOGY 

This  species  is  named  for  the  Aztec  (Nahuatl)  god  of  Spring 
and  of  flowers,  Xochipillt  in  pronoucing  the  name,  the  x has 
an  “sh”  sound. 

DISCUSSION 

Aside  from  the  variations  in  measurements  and  ratios  noted 
above,  the  females  of  this  species  are  all  very  similar  to  one 
another.  Even  the  yellowish  face  marks  appear  to  be  quite 
stable,  but  this  may  be  due  to  the  fact  that  all  are  from  the 


same  locality.  Much  the  same  is  true  of  the  males.  The  prin- 
cipal variation  involves  the  extent  of  the  shiny  area  of  the 
clypeus.  In  a few  males,  up  to  one-half  of  the  discal  area  is 
shiny,  but  generally  the  shiny  portion  is  less  extensive,  and 
may  be  limited  to  a very  narrow  band  along  the  apical  mar- 
gin. 

Most  males  possess  a narrow,  transverse  supraclypeal  mark, 
but  in  one  it  is  merely  a small  median  spot.  Other  males, 
including  the  holotype,  lack  a supraclypeal  mark.  All  males 
seen  possess  a broad  stripe  on  the  underside  of  the  scape 
which  almost  attains  the  apex  of  the  segment.  The  black 
laterobasal  marks  on  the  clypeus  are  consistently  large. 

The  combination  of  pale  ochreous  pubescence  and  bicol- 
ored abdomen  will  separate  both  sexes  of  C.  xochipHlii  from 
all  other  species.  Males  of  C.  eurypatana,  C.  heithausi,  and 
C.  xochipillii  all  have  the  basal  margin  of  the  clypeus  more 
strongly  arched  upward  in  the  center  than  at  either  side;  in 
C.fuscata  the  margin  is  evenly,  very  slightly  convex  between 
the  subantennal  sutures;  C.  dentata  is  intermediate  between 
these  two  types. 

From  C.  eurypatana,  males  of  C.  xochipillii  may  be  sep- 
arated by  the  lack  of  procoxal  spines,  the  broader  clypeus 
and  the  reduced  yellow  areas  on  the  clypeus.  In  C.  xochipillii 
the  pubescence  is  slightly  yellowish,  the  disc  of  the  second 
tergum  is  only  moderately  shiny,  and  the  punctures  of  the 
apical  zone  of  the  second  tergum  are  not  conspicuously  finer 
than  those  of  the  disc;  these  features  will  separate  C.  xo- 
chipillii from  C.  heithausi.  Males  of  C.  dentata  have  the 
punctures  of  the  clypeal  disc  distinct,  rather  than  obscured 
by  dense  tessellation,  the  disc  of  the  second  tergite  is  densely 
punctate,  and  the  ventral  tooth  of  the  metafemur  slender  and 
curved.  All  of  the  species  differ  from  C.  xochipillii  in  having 
the  abdomen  wholly  ferruginous. 

The  female  shares  with  those  of  C.  dentata,  C.  heithausi, 
and  C.  eurypatana  the  broad  secondary  plate  on  the  basitibial 
plate,  thus  differing  from  that  of  C.  fuscata.  It  is  separable 
from  all  by  the  bicolored  abdomen,  from  C.  heithausi  by  the 
ochreous  rather  than  whitish  pubescence,  from  C.  dentata 
by  the  immaculate  scape  and  shiny,  sparsely  punctate  disc 
of  the  second  tergite,  and  from  C.  eurypatana  by  the  shiny 
median  clypeal  line,  immaculate  scape,  and  much  paler  wings. 

Subgenus  Hemisiella  Moure 

Hemisiella  Moure,  1945b:407-408.  Type-species:  “ Hemi- 
siella lanipes  (Fabricius,  1775)”  = Apis  lanipes  Fabricius, 
1775;  original  designation. 

Centris  subg.  Hemisiella:  Michener,  1951:7-8. 

This  is  a large  group,  with  most  of  the  species  occurring  in 
South  America,  but  with  one  entering  the  southwestern  United 
States.  The  distinctions  between  this  subgenus  and  Hetero- 
centris  are  not  great  and  possibly  the  two  should  be  merged. 
A detailed  study  of  the  extensive  South  American  fauna  might 
resolve  this  question,  but  is  beyond  the  scope  of  this  paper. 

In  addition  to  the  characters  noted  in  the  keys  by  Michener 
(1951)  and  Snelling  (1974),  Hemisiella  females  may  be  sep- 
arated from  those  of  Heterocentris  by  the  lack  of  compressed. 


44  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


blade-like  setae  on  the  fourth  and  fifth  sterna  and  by  the 
acute  apex  of  the  secondary  pygidial  plate. 

KEY  TO  NORTH  AMERICAN  HEM  I SI  ELLA 

la.  Male,  antenna  1 3-segmented;  abdomen  with  seven  vis- 
ible terga  2 

b.  Female,  antenna  1 2-segmented;  abdomen  with  six  vis- 
ible terga  6 

2a.  Hairs  of  thoracic  dorsum  with  conspicuous  black  tips 

3 

b.  Hairs  of  thoracic  dorsum  without  conspicuous  black 

tips  4 

3a.  Large  species,  head  width  over  5.2  mm;  face  broad, 

frontal  width  greater  than  clypeocellar  distance  

vittata  Lepeletier 

b.  Smaller  species,  head  width  less  than  4.7  mm;  face 
narrower,  frontal  width  less  than  clypeocellar  distance 

'• dichrootricha  Moure 

4a.  Midline  of  clypeus  not  cariniform;  paraocular  area  with 
yellow  mark;  segments  10  and  11  of  flagellum  dark 
beneath,  contrasting  with  underside  of  segments  2-9 

5 

b.  Midline  of  clypeus  low-cariniform;  paraocular  area 
without  yellow  mark;  segments  10  and  1 1 of  flagellum 
beneath  not  contrasting  with  color  of  segments  2-9  . . 

transversa  Perez 

5a.  At  least  two  terga,  often  entire  abdomen,  ferruginous, 
apical  hairs  ferruginous;  first  flagellar  segment  less  than 

3.7  times  second  trigonoides  Lepeletier 

b.  Abdominal  terga  blackish,  apical  segments  with  white 
hairs;  first  flagellar  segment  at  least  4.0  times  second 

nitida  F.  Smith 

6a.  Hairs  of  thoracic  dorsum  conspicuously  black-tipped 

7 

b.  Hairs  of  thoracic  dorsum  not  black-tipped  8 

7a.  Clypeus  broadly  and  deeply  depressed  across  lower  part 
of  disc;  labrum  at  least  twice  wider  than  long;  larger 

species,  head  width  at  least  6.0  mm  

vittata  Lepeletier 

b.  Clypeus  protuberant;  disc  flat  or  slightly  convex;  la- 
brum no  more  than  1.5  times  wider  than  long;  smaller 

species,  head  width  less  than  5.5  mm  

dichrootricha  Moure 

8a.  Abdomen  and  legs  blackish;  scopa  black  9 

b.  Abdomen  and  middle  and  hind  legs  largely  ferruginous; 

scopa  ferruginous trigonoides  Lepeletier 

9a.  Fifth  tergum  usually  with  long,  glistening,  white  hairs; 
lower  margin  of  clypeal  marks  parallel  to  apical  margin 
of  clypeus;  smaller  species,  head  width  3. 9-4. 5 mm 

transversa  Perez 

b.  Fifth  tergum  with  brown  to  black  hairs  only;  lower 
margin  of  clypeal  marks  oblique  to  apical  margin  of 

clypeus;  larger  species,  head  width  4. 8-5. 3 mm  

nitida  F.  Smith 

Centris  (Hemisiella)  dichrootricha  (Moure) 

Hemisiella  dichrootricha  Moure,  1945b:408-409.  9. 


Centris  (Hemisiella)  dichrootricha:  Michener,  1954:143.  9 <3 
(distr.,  tax.). 

This  species  was  described  from  the  State  of  Guapore,  Brazil; 
Michener  (1954)  recorded  it  from  Panama  and  noted  char- 
acteristics of  the  previously  undescribed  male.  I have  seen 
many  specimens  from  Panama  and  a few  from  Costa  Rica. 
The  species  is  easily  recognized  by  the  characteristics  cited 
in  the  key. 

Centris  ( Hemisiella ) nitida  F.  Smith 

Centris  nitida  F.  Smith,  1874:368.  9. 

Centris  confinis  Perez,  1905:40.  9.  NEW  SYNONYMY. 

Perez  (1905)  described  C.  confinis  from  “Mexique?”  The 
type  specimen,  a female,  is  in  the  Museum  National  d'His- 
toire  Naturelle,  Paris,  and  has  been  examined  by  me.  Al- 
though in  poor  condition,  it  is  unquestionably  a specimen  of 
C.  nitida.  The  records  from  southern  Arizona  cited  by  Hurd 
( 1 979)  as  C.  confinis  are  based  on  specimens  of  C.  transversa. 

This  is  a common  species  through  Central  America,  ex- 
tending into  northern  South  America;  the  type  locality  is 
Honduras,  without  more  precise  locality.  I have  seen  Central 
American  material  of  C.  nitida  from  Mexico,  Belize,  Gua- 
temala, Honduras,  El  Salvador,  and  Costa  Rica.  Although  I 
have  seen  no  records  from  Panama,  C.  nitida  must  be  present 
there,  as  it  is  present  in  South  America  (Colombia,  Ecuador). 

Centris  ( Hemisiella ) transversa  Perez 

Centris  transversa  Perez,  1905:39.  9 6. 

Hemisiella  transversa:  Moure,  1945b:408. 

Centris  (Melanocentris)  ruae  Cockerell,  1949:474-475.  9. 
NEW  SYNONYMY. 

Centris  (Hemiesiella)  transversa:  Snell  mg,  1966:26-27  (distr.). 
Centris  (Hemisiella)  confinis:  Hurd,  1979:2175  (misidenti- 
fication). 

This  primarily  Mexican  species  is  found  also  in  Guatemala 
and  Honduras.  Although  it  has  been  suggested  that  this  is  a 
synonym  of  C.  nitida  (see  Lutz  and  Cockerell,  1920:560),  the 
two  are  distinct  from  one  another.  In  addition  to  differences 
noted  in  the  key,  females  of  C.  transversa  have  the  median 
impunctate  line  of  the  clypeus  distinctly  raised,  the  flagellum 
is  uniformly  dark,  the  dorsal  thoracic  hairs  are  brownish  and 
the  hairs  at  the  sides  of  the  fourth  and  fifth  sternites  are 
whitish.  In  C.  nitida,  the  impunctate  median  line  is  not  raised 
or,  if  a little  elevated,  the  raised  area  is  broadly  rounded 
rather  than  narrow  and  sharply  defined;  the  underside  of  the 
flagellum  is  lighter  in  color  than  the  upper  side;  the  dorsal 
thoracic  hairs  are  yellowish;  the  hairs  at  the  sides  of  the  fourth 
and  fifth  sternites  are  uniformly  dark. 

Males  of  C.  transversa  possess  a somewhat  coniform  ven- 
tral process  on  the  metatrochanter,  a prominent  ventral  pro- 
cess near  the  base  of  the  metafemur,  and  the  pubescence  of 
the  fifth  and  sixth  sternites  is  mostly  pale.  The  ventral  process 
of  the  metatrochanter  of  C.  nitida  is  a depressed,  narrow 
apical  spine  and,  in  that  species,  the  ventral  surface  of  the 
metafemur  lacks  a process  and  the  pubescence  of  the  fourth 
to  sixth  sterna  is  dark. 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  45 


The  type  of  C.  ( Melanocentris ) ruae  is  in  the  USNM  (No. 
58880)  and  is  from  Zamorano,  Honduras.  I have  examined 
the  specimen  and  it  is  a normal  specimen  of  C.  transversa, 
differing  from  Mexican  specimens  only  in  the  reduction  in 
the  number  of  white  hairs  on  the  apical  terga.  In  spite  of 
Cockerell’s  statement  to  the  contrary,  the  size  is  normal  and 
the  clypeal  and  labral  markings  are  well  within  the  range  for 
this  species.  The  type  agrees  with  other  material  from  Hon- 
duras. 

Hurd  (1979)  reported  C.  (Hemisiella)  confinis  from  Ari- 
zona (Patagonia  and  Tumacacori)  on  flowers  of  Parkinsonia 
aculeata.  The  specimens  on  which  this  report  was  based  are 
in  UCB  and  I have  examined  them.  They  are  C.  transversa, 
which  I had  previously  (1966)  recorded  from  Arizona.  At 
that  time  I conjectured  that  C.  transversa  was  possibly  ad- 
ventive  in  Arizona,  a view  I no  longer  hold.  Additional  ma- 
terial now  shows  the  distribution  of  C.  transversa  to  extend 
north  along  the  western  flanks  of  the  Sierra  Madre  Occidental 
through  Sonora  to  southern  Arizona. 


Centris  ( Hemisiella ) trigonoides  Lepeletier 

Centris  trigonoides  Lepeletier,  1841:167.  <5. 

Centris  dentipes  F.  Smith,  1874:366.  <5.  NEW  SYNONYMY. 
Centris  hoplopoda  Moure,  1943:160.  <5. 

Centris  rufomaculata  Cockerell,  1949:476.  <5.  NEW  SYN- 
ONYMY. 

Centris  ( Rhodocentris ) lanipes  subtarsata  Cockerell,  1949: 
476-477.  6 2.  NEW  SYNONYMY. 

Centris  ( Hemisiella ) hoplopoda:  Michener,  1954:142-143 
(distr.,  tax.). 

Centris  (Hemisiella)  trigonoides  subtarsata:  Snelling,  1966: 
25-26  (distr.,  tax.). 

Centris  trigonoides  is  a common  species,  ranging  from  Mex- 
ico to  Argentina.  Throughout  this  range  it  is  subject  to  much 
variation  and  some  localized  phenotypes  have  been  named; 
presumably  there  are  additional  synonyms  to  be  recognized 
among  the  many  names  applied  to  South  American  forms 
of  Hemisiella.  The  entire  complex  of  forms  in  this  difficult 
group  will  have  to  be  examined. 

In  an  earlier  paper  (Snelling,  1966),  I attempted  to  justify 
recognition  of  C.  lanipes  subtarsata  as  a Central  American 
subspecies  of  C.  trigonoides.  Subsequent  study  of  several 
hundred  additional  specimens  convinced  me  of  the  futility 
of  that  effort. 

The  type  male  of  C.  dentipes  (BMNH  17B.919)  has  been 
examined  and  agrees  with  the  current  concept  of  C.  trigo- 
noides. The  type  male  of  C.  rufomaculata  (USNM  58883)  is 
merely  an  individual  with  abdomen  mostly  dark  brownish. 

Females  of  C.  trigonoides  consistently  have  the  entire  ab- 
domen ferruginous  in  Central  American  samples,  and  cannot 
be  confused  with  any  other  species  in  our  area.  The  abdomen 
in  males  varies  from  wholly  ferruginous  to  dark  reddish  brown 
on  the  basal  three  or  four  tergites.  The  metatrochanter  has 
a prominent,  thick,  ventral  spine,  the  ventral  ridge  of  the 


metafemur  is  high  and  cariniform,  and  the  ventral  pubes- 
cence of  the  abdomen  is  yellowish  to  reddish. 

Centris  ( Hemisiella ) vittata  Lepeletier 

Centris  vittata  Lepeletier,  1841:168.  <3  2. 

Centris  montezuma  Cresson,  1879:213.  2 3. 

Centris  breviceps  Friese,  1899:44.  $ 2. 

Centris  Friesei  Crawford,  1906:158.  2.  Preoccupied. 

Centris  Costaricensis  Crawford,  1907:21.  New  name  for  C. 

friesei  Crawford,  1906,  not  C.  friesei  Ducke,  1902. 
Centris  costaricensis  var.  erubescens  Friese,  1 925:30.  2.  NEW 
SYNONYMY. 

Both  sexes  of  C.  vittata  are  easily  recognized  by  the  large  size 
(length  over  20  mm),  black-tipped  thoracic  hairs,  and  the 
transversely  depressed  clypeal  disc.  The  posteroventral  mar- 
gin of  the  male  metafemur  is  sharply  angled,  but  is  not  a 
cariniform  ridge  and  the  ventral  spine  of  the  metatrochanter 
is  reduced  to  an  inconspicuous,  obtuse  tubercle. 

This  is  a widespread  species  through  South  America,  and 
in  Central  America.  1 have  seen  specimens  from  Mexico, 
Costa  Rica,  Panama,  and  Honduras.  Friese's  C.  costaricensis 
var.  erubescens,  described  from  Costa  Rica,  is  a minor  vari- 
ant, not  worthy  of  recognition. 

Subgenus  Heterocentris  Cockerell 

Gundlachia  Cresson,  1865:195.  Type-species:  Centris  ? cor- 
nuta  Cresson,  1865;  type  by  monotypy.  Preoccupied. 
Heterocentris  Cockerell,  1899:14.  Type-species:  Centris  ? 
cornuta  Cresson,  1865;  autobasic.  New  name  for  Gund- 
lachia Cresson,  1865,  not  Gundlachia  Pfeiffer,  1850  (Mol- 
lusca),  not  Gundlachia  Herrich-Schaeffer,  1866  (Insecta, 
Lepidoptera). 

Centris  subg.  Rhodocentris  Friese,  1900b:  244.  Type-species: 
C.  dijformis  F.  Smith,  1854;  designated  by  Sandhouse, 
1943. 

Since  Cresson’s  generic  name  Gundlachia  was  preoccupied, 
Cockerell  (1899)  proposed  Heterocentris  as  a replacement 
name;  the  type-species  for  Heterocentris  automatically  is  C. 
? cornuta  which  Cockerell  (1906)  later  considered  to  be  the 
same  as  C.  dijformis.  There  is  no  evidence  that  he  actually 
saw  the  type  of  C.  cornuta,  now  in  the  Gundlach  collection 
of  the  Academia  de  Ciencias  in  Havana,  Cuba.  Indeed,  there 
is  reason  to  suppose  that  he  did  not,  for  the  original  descrip- 
tions of  C.  cornuta  and  C.  dijformis  do  not  agree  in  several 
important  peculiarities  of  head  structure.  Thus,  Cresson  de- 
scribed the  mandible  of  C.  cornuta  as  “very  long,  narrow 
and  cleft  at  tip,  shining  black  . . .”  as  opposed  to  Smith’s 
“.  . . mandibles  large,  very  broad  at  their  base,  and  armed 
above  with  a stout  tooth,  their  apex  bidentate,  having  a lon- 
gitudinal pale  testaceus  stripe”  (italics  mine).  Of  the  clypeus, 
Cresson  stated:  “clypeus  short,  very  transverse,  emarginate 
on  each  side,  with  a large,  very  prominent,  incurved,  subacute 
tooth  on  the  middle,  pale  yellowish  white,  the  anterior  and 
posterior  margin  and  the  tooth  except  its  lateral  base,  black” 
(italics  mine);  the  labrum  is  said  to  possess  a “long  slender. 


46  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


porrect,  subacute  spine.”  According  to  Smith,  the  clypeus  of 
C.  dijformis  is  “short,  transverse,  elevated,  its  anterior  por- 
tion vertical.”  There  is  no  mention  of  processes  of  any  sort 
on  either  clypeus  or  labrum.  I think  it  as  unlikely  that  Cresson 
would  have  overlooked  the  very  prominent  mandibular  pro- 
cess as  that  Smith  would  have  failed  to  mention  such  a con- 
spicuous anomaly  (in  this  genus)  as  a spinose  clypeus.  It  is 
my  opinion  that  C.  cornuta  and  C.  difformis  are  very  different 
entities  and  that  C.  cornuta  must  be  properly  considered  to 
be  the  type-species  of  Heterocentris. 

This  possibly  has  unfortunate  ramifications  since  the  iden- 
tity of  C.  cornuta  is  problematic.  However,  since  the  only 
forms  of  Centris  with  unusual  modifications  of  mandible, 
labrum,  and  clypeus  all  belong  to  the  accepted  interpretation 
of  Heterocentris,  the  question  is  probably  moot. 

Just  as  the  type-species  of  Heterocentris  should  be  clearly 
restricted  to  C.  cornuta,  so,  too,  should  the  type-species  of 
Rhodocentris  be  restricted  to  C.  dijformis.  Rhodocentris  was 
described  as  a new  subgenus  of  Centris,  not  as  a replacement 
name  for  Gundlachia.  It  is  clear  that  Sandhouse  considered 
C.  dijformis  to  be  the  proper  name  for  the  species  which  she 
selected  as  type.  It  is  unclear  why  she  chose  to  cite  the  type- 
species  as  “( Centris  cornuta  Cresson,  1 865)  = Centris  diffor- 
mis  F.  Smith,  1854,”  unless  it  was  to  assure  that  Rhodocentris 
was  an  automatic  junior  synonym  of  Heterocentris  through 
isogenotypy.  Both  specific  names  were  available,  since  both 
were  originally  included  in  Rhodocentris  by  Friese.  Since  it 
is  clear  that  Sandhouse  was  of  the  opinion  that  C.  difformis 
was  the  correct  name  for  the  taxon  chosen  as  type-species  I 
think  it  best  to  consider  that  name  to  be  the  type;  this  would 
eliminate  the  ambiguity  of  having  two  names  involved  as 
possible  type-species. 

Heterocentris,  together  with  Hemisiella  and  Trachina,  is 
part  of  a complex  recognized  by  possessing  three-segmented 
maxillary  palps  in  both  sexes  and  the  male  with  a carina 
along  the  posterior  margin  of  the  metabasitarsus;  this  carina 
usually  terminates  in  a prominent  tooth-like  process.  Males 
of  Heterocentris  differ  from  those  of  both  Trachina  and 
Hemisiella  in  the  form  of  the  dorsal  face  of  the  first  abdom- 
inal tergite:  at  the  extreme  side,  the  dorsal  face  is  extended 
caudad  (most  strongly  so  in  C.  labrosa ) and  there  is  a con- 
spicuous patch  of  erect,  plumose,  dark  setae  at  the  side  of 
the  segment,  their  apices  abruptly  bent  and  llattened.  This 
is  a feature  unique  in  the  genus.  Additionally,  the  middle 
mandibular  tooth  is  smaller  and  nearer  to  the  inner  tooth 
than  to  the  apical  tooth. 

In  females  of  Heterocentris  the  upper  inner  mandibular 
carina  is  elevated  near  the  base  and  the  labrum  is  large,  with 
the  disc  depressed  and  the  apical  margin  more  or  less  flange- 
like and  with  a pair  of  dentiform  submedian  processes.  In 
both  sexes  the  apicolateral  angle  of  the  clypeus  is  contiguous 
with  the  eyes,  or  nearly  so.  Females  are  additionally  char- 
acterized by  the  distinct,  abruptly  truncate  secondary  pygidial 
plate  and  by  the  presence  of  large,  flattened,  spiniform  setae 
near  the  apical  margins  of  the  fourth  and  fifth  abdominal 
sterna. 

The  few  Central  American  Heterocentris  may  be  separated 


by  the  following  key.  Since  no  males  of  C.  difformis  are 
known  to  me,  I am  unable  to  include  this  sex  in  the  key. 

KEY  TO  CENTRAL  AMERICAN  HETEROCENTRIS 

la.  Male,  antenna  13-segmented  and  basitibial  plate  absent 

2 

b.  Female,  antenna  12-segmented  and  basitibial  plate  pres- 
ent   5 

2a.  Postgradular  area  of  second  tergum  about  four  times  as 

long  as  pregradular  area  3 

b.  Postgradular  area  of  second  tergum  shorter  than  pre- 
gradular area labrosa  Friese 

3a.  Mandible  tridentate 4 

b.  Mandible  bidentate  bicornuta  Mocsary 

4a.  Clypeal  disc  smooth  and  shiny  between  punctures;  apical 
margin  of  labrum  transverse  or  very  weakly  concave; 
lower  corner  of  pronotum  with  a few  long,  simple,  red- 
dish or  yellowish  setae  among  plumose  hairs  

analis  (Fabricius) 

b.  Clypeal  disc  roughened  and  slightly  shiny  between  punc- 
tures; apical  margin  of  labrum  with  distinct  median 
emargination;  lower  corner  of  pronotum  with  plumose 

hairs  only  C.  (Heterocentris)  species 

5a.  Clypeus  without  lateral  cornuti;  labrum  broader  than 

long;  mandible  tridentate  5 

b.  Clypeus  with  pair  of  long,  slender  cornuti;  labrum  longer 
than  broad;  mandible  bidentate  bicornuta  Mocsary 
6a.  Clypeus  no  more  than  1 .8  times  wider  than  long  in  mid- 
dle, with  disc  flattened;  mandible  without  long  subbasal 

process  on  outer  face  7 

b.  Clypeus  very  short,  more  than  twice  wider  than  long, 
transversely  elevated;  mandible  with  large,  tooth-like 
subbasal  process  on  outer  face  ....  difformis  F.  Smith 
7a.  Lower,  lateral  angle  of  pronotum  with  a cluster  of  long, 
yellowish  or  reddish,  simple  setae  as  well  as  plumose 
hairs;  juncture  of  anterior  and  lateral  faces  of  mesepi- 
sternum  without  carina,  rounded;  hairs  of  thoracic  dor- 
sum rarely  dark-tipped analis  (Fabricius) 

b.  Lower,  lateral  angle  of  pronotum  with  long,  plumose 
hairs  only;  juncture  of  anterior  and  lateral  faces  of  mes- 
episternum  with  a short,  lamelliform  carina;  hairs  of 
thoracic  dorsum  always  dark-tipped  . . . labrosa  Friese 

Centris  ( Heterocentris ) analis  (Fabricius) 

Anthophora  analis  Fabricius,  1804:375.  2. 

Centris  totonaca  Cresson,  1879:213.  2. 

Centris  otomita  Cresson,  1879:214.  <3. 

Centris  minuta  Mocsary,  1899:254.  <3. 

Centris  labrosa  var.  simplex  Friese,  1 899:44.  2. 

Centris  (Melanocentris)  durantae  Cockerell,  1949:474.  <3. 
Centris  (Melanocentris)  petreae  Cockerell,  1949:475.  <3. 
Centris  (Melanocentris)  petreae  var.  rufopicta  Cockerell,  1 949: 
475.  <3. 

Centris  (Heterocentris)  totonaca:  Michener.  1954:140  (syn.). 
Centris  (Heterocentris)  analis:  Moure,  1 960b:  1 32-133  (syn., 
notes  on  type). 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  47 


Moure  (1960b)  gives  very  complete  literature  citations  of 
this  common  species  which  ranges  from  Mexico  to  Brazil. 
The  presence  of  long,  simple,  ferruginous  setae  on  the  lower 
corner  of  the  pronotum  is  diagnostic  for  the  female.  The  male 
has  similar  setae,  but  they  are  less  numerous  than  in  the 
female  and  are  sometimes  difficult  to  see  among  the  more 
numerous  plumose  hairs. 

Centris  (Heterocentris)  bicornuta  Mocsary 

Centris  bicornuta  Mocsary,  1899:254.  9. 

Heterocentris  bicornuta:  Moure,  1945b:502. 

Centris  (Heterocentris)  bicornuta:  Michener,  1951:6,  7. 

Although  widely  distributed,  C.  bicornuta  does  not  appear 
to  be  a common  species,  though  males  are  sometimes  locally 
abundant.  The  bicornute  clypeus  of  the  female  is  diagnostic 
for  that  sex.  Males  may  be  easily  separated  from  those  of  C. 
analis  and  C.  labrosa  by  the  bidentate  mandible.  Since  I have 
seen  no  males  of  C.  difformis,  which  presumably  has  biden- 
tate mandibles,  I am  uncertain  how  to  distinguish  that  species 
from  C.  bicornuta.  Presumably,  however,  the  males  of  C. 
difformis  will  be  more  than  1 5 mm  long  and  the  hairs  of  the 
thoracic  dorsum  will  be  black-tipped.  Males  of  C.  bicornuta 
are  not  more  than  about  10  mm  long  and  the  hairs  of  the 
thoracic  dorsum  are  not  black-tipped. 

Specimens  of  C.  bicornuta  have  been  seen  from  Mexico, 
Guatemala,  Costa  Rica,  and  Panama,  as  well  as  from  South 
America  (Brazil  and  Guyana).  The  species  was  described 
from  Brazil. 

Centris  ( Heterocentris ) difformis  F.  Smith 

Centris  difformis  F.  Smith,  1854:374.  9. 

Centris  difformis:  Crawford,  1906:158.  9. 

Heterocentris  difformis:  Moure,  1945b:  402,  403. 

Centris  ( Heterocentris ) difformis:  Michener,  1951:  6,  7. 

As  discussed  above,  1 do  not  believe  that  C.  cornuta,  de- 
scribed from  Cuba,  is  a synonym  of  C.  difformis.  The  latter 
species  was  originally  described  from  Brazil  and  seems  to  be 
rare  in  collections.  According  to  Cresson  ( 1879)  C.  difformis 
occurs  in  Mexico,  but  I have  seen  no  specimens  from  there. 
Crawford  (1906)  recorded  a female  from  Pozo  Azul,  Costa 
Rica,  as  C.  difformis.  I have  examined  the  specimen  and 
agree  with  Crawford's  identification. 

No  males  have  been  seen.  Presumably  the  mandibles  are 
bidentate,  as  in  C.  bicornuta  (Moure,  1945b.  noted  that  C. 
minuta  and  C.  labrosa  differed  from  his  characterization  of 
Heterocentris  mandibles  as  bidentate  with  the  statement  that 
the  mandible  is  falsely  tridentate  in  these  two  species).  Based 
on  the  females,  males  of  C.  difformis  should  be  conspicuously 
larger  than  those  of  C.  bicornuta  and  the  hairs  of  the  thoracic 
dorsum  should  have  blackish  apices. 

NEW  RECORDS 

PANAMA,  CANAL  ZONE:  399,  Barro  Colorado  Island.  27 
Apr.  1980  (K.E.  Steiner;  UCD),  on  Byrsonima  crassifolia. 

Centris  ( Heterocentris ) labrosa  Friese 

Centris  labrosa  Friese,  1899:44.  9 (not  3). 


Centris  tarsata:  Schwarz,  1934:13.  Misidentification. 
Heterocentris  labrosa:  Moure,  1945b:402. 

Centris  (Rhodocentris)  triangulifera  Cockerell,  1949:  477.  9. 
NEW  SYNONYMY. 

Centris  (Heterocentris)  labrosa:  Michener,  1954:104  (var., 
distr.). 

This  is  a moderately  common  species  which  ranges  from 
Mexico  to  Brazil.  I have  seen  specimens  from  throughout 
Central  America.  It  should  be  noted  that  the  specimen  re- 
corded by  Schwarz  (1934)  from  Barro  Colorado  Island,  Pan- 
ama, as  C.  tarsata  F.  Smith,  is  actually  C.  labrosa.  The  type 
of  C.  triangulifera.  in  the  USNM  (No.  58885),  has  been 
examined  and  is  a normal  specimen  of  C.  labrosa. 

The  unusually  long  pregradular  area  of  the  males  is  im- 
mediately diagnostic  for  this  sex  of  C.  labrosa.  Females  are 
likewise  immediately  recognizable  by  the  presence  of  a dis- 
tinct carina  separating  the  anterior  and  lateral  faces  on  the 
lower  half  of  the  mesepisternum;  in  all  other  species,  the 
juncture  of  the  two  surfaces  is  rounded. 

Centris  ( Heterocentris ) species 

A few  males  from  Panama  (Canal  Zone  and  Panama  Prov- 
ince) cannot  be  assigned  to  any  of  the  above  species.  It  seems 
unlikely  that  they  are  males  of  C.  difformis  for  they  are  much 
smaller  (less  than  15  mm  long)  than  the  one  female  of  C. 
difformis  (about  20  mm  long)  I have  seen;  the  two  sexes  tend 
to  be  approximately  equal  in  size  in  this  genus. 

They  are  very  similar  to  males  of  C.  analis.  but  lack  the 
simple  setae  on  the  lower  lateral  angle  of  the  pronotum,  the 
clypeus  is  distinctly  dull  and  roughened  between  the  punc- 
tures, and  the  mandible  and  scape  are  immaculate  or  with 
traces  of  yellowish  markings.  Possibly  these  represent  an  un- 
described species,  but  the  available  material  is  too  limited 
for  any  decision  as  to  their  identity. 

Genus  Ptilotopus  Klug 

Ptilotopus  Klug,  1810:31.  Type-species:  Ptilotopus  ameri- 
canus  Klug,  1810;  monobasic. 

Centris  subg.  Ptilotopus:  Michener,  1951:10.  Snelling,  1974: 
2,  3. 

In  recent  years  Ptilotopus  has  usually  been  treated  as  a sub- 
genus of  Centris  characterized  by  the  bilobate  scutellum  with 
defined  bare  areas,  the  prominent  hypoepimeral  tubercle,  the 
lack  of  a secondary  basitibial  plate  in  the  female  and  the  male 
genitalia  without  giant  branched  setae  (Michener,  1951; 
Snelling.  1974).  The  largest  and  most  spectacular  centridines 
are  included  in  Ptilotopus. 

Although  in  general  habitus  the  species  of  Ptilotopus  are 
similar  to  those  of  Centris  and  have  Centris- like  wing  ve- 
nation, I am  now  of  the  opinion  that  Ptilotopus  should  be 
removed  from  Centris.  There  are  a number  of  features  which 
are  not  shared  with  any  of  the  groups  presently  assigned  to 
Centris.  Both  sexes  of  Ptilotopus  possess  long,  black  flagel- 
liform  setae  on  the  occipital  margin;  these  setae  project  well 
beyond  the  occipital  hair  fringe,  although  they  are  not  as 
spectacularly  developed  as  in  many  Epicharis. 


48  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


Contributions  in  Science,  Number  347 


Sneiling:  American  Centridini  49 


The  metatibia  of  Ptilotopus  females  has  a well-defined  ba- 
sitibial  plate.  There  is,  however,  no  secondary  plate;  instead, 
the  disc  of  the  basitibial  plate  is  moderately  depressed,  the 
depression  filled  with  a dense  mat  of  short,  fine,  subappressed 
hairs.  In  Centris  and  Epicharis  females,  a secondary  plate  is 
present  and  usually  well  defined.  When  it  is  not  distinct  the 
plate  has  a discal  convexity  and  is  glabrous. 

Females  of  Ptilotopus  have  a distinct  tubercle  on  either 
side  of  the  midline  of  the  mesosternum,  anterior  to  the  me- 
socoxa.  As  a rule  the  tubercles  are  hidden  in  an  exceptionally 
dense  tuft  of  short,  stiffhairs.  Although  mesosternal  tubercles 
are  known  for  some  Epicharis  males,  they  are  not  known  to 
be  present  in  the  females  of  either  Centris  or  Epicharis. 

A particularly  unusual  feature  is  to  be  seen  in  the  structure 
of  the  basal  areas  of  the  second  and  third  abdominal  tergites 
of  the  females.  In  both  Centris  and  Epicharis  the  gradulus 
of  each  segment  marks  off  a very  narrow  basal  area  across 
most  of  the  breadth  of  the  segment;  laterad.  it  is  either  eva- 
nescent or  deflected  distad.  In  Ptilotopus  females  the  gradulus 
of  the  second  and  third  tergites  is  strong  and,  in  the  middle, 
is  directed  distad  as  a more  or  less  triangular  incursion  onto 
the  disc  of  the  segment.  The  area  on  either  side  of  this  tri- 
angular extension  is  depressed  and  filled  with  a compact  mat 
of  very  short,  erect,  plumose  hairs.  This  unusual  feature  is 
not  known  to  occur  in  Centris  and  in  Epicharis  is  known 
only  in  the  subgenus  Epicharitides. 

Thus,  although  Ptilotopus  is  Centris-Yxkt  in  many  features, 
particularly  in  general  body  form  and  wing  venation,  there 
are  features,  such  as  the  presence  of  the  flagelliform  occipital 
setae,  presence  of  mesosternal  tubercles  in  the  female  and 
the  modified  structure  of  the  female  second  and  third  ab- 
dominal terga,  which  will  readily  separate  Ptilotopus  from 
Centris.  The  short  marginal  cell  of  the  forewing  will  readily 
separate  Ptilotopus  from  Epicharis,  as  will  the  mesosternal 
tubercles  and  lack  of  a secondary  basitibial  plate  in  the  fe- 
male. Additionally,  in  Ptilotopus,  the  hypoepimeron  has  a 
prominent  tubercle  and  the  scutellum  has  a conspicuous, 
flattened,  bare  lobe  on  either  side  of  the  middle.  The  genitalic 
structures  of  Ptilotopus  are  very  distinctly  Centra-like,  with- 
out the  unusual  modifications  seen  in  Epicharis. 


Ptilotopus  zouata  (Mocsary) 

Figures  59-62 

Centris  zonata  Mocsary,  1899:251.  9. 

Centris  pandora  Friese,  1900b:241,  269  (new  name  for  C. 
zonata  Mocsary,  not  C.  zonata  F.  Smith,  1 854,  now  placed 
in  Epicharis). 

This  species,  the  only  known  North  American  Ptilotopus, 
was  described  from  Chiriqui  Province,  Panama;  no  addi- 
tional material  has  been  recorded  since  the  original  descrip- 
tion. 

The  females  have  the  pubescence  of  the  head,  thorax,  and 
legs  bright  ferruginous.  Except  for  a broad  band  of  short, 
erect  yellow  pubescence  across  the  second  tergite,  the  ab- 


domen is  black  pubescent.  The  male  pubescence  is  all  black, 
except  for  a yellow  band  across  the  second  tergite. 

NEW  RECORDS 

PANAMA,  PANAMA:  1499,  Arraijan,  16  Oct.  1980(D.  Rou- 
bik;  LACM,  ROUB).  VER.4GUAS:  13,  El  Maria,  Coibu  Is- 
land, 22  Oct.  1979  (D.  Roubik;  LACM).  CANAL  ZONE:  12, 
Gatun,  3 Nov.  1977  (K.E.  Steiner;  UCD),  on  Stigmaphyllon 
hypargyreum. 

Genus  Epicharis  Klug 

Epicharis  is  an  exclusively  Neotropical  genus  of  medium-  to 
large-sized  bees,  often  quite  colorful,  allied  to  Centris.  Friese 
( 1 900b)  monographed  these  bees,  as  a subgenus  of  Centris, 
but  most  subsequent  authors  have  treated  Epicharis  as  a 
separate  genus.  Since  Friese’s  monograph  most  work  in  this 
genus  has  consisted  ofdescriptions  of  new  species  and  variant 
forms.  Moure  ( 1945a),  however,  divided  Epicharis  into  nine 
genera,  for  which  he  proposed  the  subtribe  Epicharitina  with- 
in within  the  Centridini.  Michener  ( 1954)  recognized  Moure’s 
genera  as  subgenera  within  the  single  genus  Epicharis  and 
subsequent  authors  have  been  in  accord  with  this  interpre- 
tation. 

Epicharis  was  described  by  Klug  ( 1 807),  but  a type-species 
was  not  designated  until  Latreille  (1810)  fixed  Apis  rustica 
Olivier,  1789,  as  the  type-species;  A.  rustica  was  not  one  of 
the  originally  included  species  and,  hence,  is  not  available. 
Lutz  and  Cockerell  (1920)  selected  Centris  umbraculata  Fa- 
bricius,  1804,  one  of  the  originally  included  species,  as  the 
type-species.  Moure  ( 1 945a,  1 960b)  held  that  the  designation 
made  by  Latreille  is  valid  since  Apis  hirtipes  Fabricius,  1 793, 
one  of  the  originally  included  names,  is  a synonym  of  A. 
rustica. 

Sandhouse  (1943)  accepted  the  designation  of  Lutz  and 
Cockerell,  as  did  Michener  (1954)  when  he  named  Epicha- 
rana  to  replace  Epicharis  (s.s.)  of  Moure  ( 1945a)  and  placed 
Xanthepicharis  Moure  in  synonymy  with  true  Epicharis  (s.s.). 
It  is  my  understanding  of  the  ICZN  Code  that  even  though 
A.  hirtipes  ultimately  proved  to  be  a synonym  of  A.  rustica, 
this  does  not  validate  the  designation  made  by  Latreille,  since 
Latreille  was  not  aware  that  the  two  names  applied  to  the 
same  species.  There  is,  additionally,  the  fact  that  A.  hirtipes 
can  never  be  anything  other  than  a subjective  synonym  of  A. 
rustica ; the  synonym  is  a generally  accepted  opinion  which 
has  the  potential  for  being  incorrect.  Stability  would  be  best 
served  by  accepting  the  security  of  the  designation  of  Lutz 
and  Cockerell. 

Of  the  nine  subgenera  recognized  by  Moure  (1945a),  five 
are  known  to  be  present  in  North  America;  Hoplepicharis 
Moure,  1945a;  Epicharana  Michener,  1945  (=“ Epicharis” 
of  Moure,  1945a);  Epicharoides  Radoszkowski,  1884;  Pa- 
repicharis  Moure,  1945a;  and  Epicharitides  Moure,  1945a. 
The  remaining  four  subgenera,  including  Epicharis  proper 
(=Xanthepicharis  Moure,  1 945a),  are  limited  to  South  Amer- 
ica: Anepicharis,  Triepicharis,  and  Cyphepicharis,  all  Moure, 
1945a. 


50  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


The  taxonomy  of  Epicharis  is  difficult.  These  bees  are 
generally  black  or  blackish  and  both  sexes  usually  have  con- 
spicuous patterns  of  white  or  yellow  and/or  ferruginous 
markings  on  various  parts  of  the  body.  The  species  within  a 
subgenus  tend  to  be  morphologically  very  similar  and  dif- 
ferences often  are  subtle.  Previous  descriptive  work  has  tend- 
ed to  emphasize  differences  in  maculations.  This,  coupled 
with  a scarcity  of  specimens,  has  resulted  in  a confused  sit- 
uation. I expect  a reduction  in  the  number  of  species  of 
Epicharis  as  more  material  becomes  available  and  the  species 
better  known.  I have  proposed  some  new  synonymy  below 
and  pointed  out  cases  where  I suspect  further  synonymy  will 
be  in  order.  Two  new  species  are  described,  based  on  mor- 
phological characteristics. 

The  following  key  to  the  subgenera  of  Epicharis  is  modified 
from  that  of  Moure  (1945a). 

KEY  TO  SUBGENERA  OF  EPICHARIS 

la.  Female  with  secondary  basitibial  plate;  male  with  py- 
gidial  plate  obsolete,  its  margins  more  or  less  coextensive 
with  margins  of  seventh  tergite,  apex  thin  and  bilobate 

or  bidentate 2 

b.  Female  without  secondary  basitibial  plate;  male  with 
distinct,  sharply  marginate  pygidial  plate,  no  more  than 
one-half  as  wide  as  seventh  tergite,  its  apex  narrowly 

truncate  or  rounded  4 

2a.  Lateral  margins  of  female  pygidial  plate  distinctly  con- 
cave in  dorsal  view,  apex  broadly  truncate  (Fig.  63); 
female  metatibia  no  longer  than  metabasitarsus;  male 
without  scopa-like  hairs  on  hind  legs  and  metabasitarsus 
with  longitudinal  keel  on  anterior  margin  which  termi- 
nates in  spiniform  process  at  apex,  or  without  keel.  . 3 
b.  Lateral  margins  of  female  pygidial  plate  nearly  straight, 
apex  narrowly  truncate  (Fig.  64);  female  metatibia  longer 
than  metabasitarsus;  hind  legs  of  male  with  long,  scopa- 
like  hairs  and  metabasitarsus  with  prominent  tooth  at 

basal  one-third  Hoplepicharis 

3a.  Female:  frontal  carina  ending  more  than  diameter  of 
anterior  ocellus  in  front  of  that  ocellus;  pygidial  trun- 
cation narrower  than  diameter  of  anterior  ocellus;  disc 
of  fifth  tergite  with  very  short  hairs  mostly  simple  or 
barbulate.  Male:  mesosternal  tubercles  absent;  meta- 
basitarsus without  carinate  ridge  on  anterior  margin; 
metatrochanter  and  metafemur  without  ventral  patch  of 

short,  dark  plumose  setae  Epicharis,  s.s. 

b.  Female:  distance  between  frontal  carina  and  anterior 
ocellus  less  than  diameter  of  anterior  ocellus;  pygidial 
truncation  broader  than  diameter  of  anterior  ocellus; 
disc  of  fifth  tergite  with  very  short  hairs  distinctly  plu- 
mose. Male:  mesosternal  tubercles  present;  metabasi- 
tarsus with  carinate  ridge  on  anterior  margin,  terminat- 
ing in  tooth-like  process  (Fig.  65);  metatrochanter  and 
metafemur  with  ventral  patch  of  short,  dark,  plumose 

setae  Epicharana 

4a.  First  flagellar  segment  short,  in  female  usually  no  longer 
than  combined  second  and  third  but  always  shorter  than 
following  three  combined,  and  in  male  shorter  than  scape; 


posterior  margin  of  dorsal  face  of  scutell  um  not,  or  weak- 
ly, impressed  in  middle  (if  deeply  impressed,  maxillary 

palp  three-segmented) 5 

b.  First  flagellar  segment  of  female  equal  to  length  of  next 
three  segments  combined,  of  male  longer  than  scape  and 
longer  than  following  two  segments  combined;  posterior 
margin  of  dorsal  face  of  scutellum  deeply  impressed  in 
middle;  maxillary  palp  two-segmented  . Parepicharis 
5a.  Lateral  ridges  of  clypeal  disc  absent  or  weakly  evident 
only  on  basal  one-half  or  less;  male  metabasitarsus  with 
carinate  ridge  on  anterior  margin,  terminating  in  large 

tooth  at  about  midlength  6 

b.  Lateral  ridges  of  clypeal  disc  strong  and  sharply  defined 
to  near  apical  margin;  male  metabasitarsus  without  car- 
inate ridge  or  tooth  on  anterior  margin  7 

6a.  Maxillary  palp  two-segmented;  fiagelliform  occipital  se- 
tae distinct  and  extending  beyond  anterior  margin  of 
mesoscutum;  ocellocular  distance  of  male  less  than  di- 
ameter of  lateral  ocellus  in  dorsal  view  . . Anepicharis 
b.  Maxillary  palp  three-segmented;  fiagelliform  occipital 
setae  short,  not  projecting  beyond  occipital  hairs  and  not 
reaching  anterior  margin  of  mesoscutum;  ocellocular 
distance  of  male  equal  to  diameter  of  lateral  ocellus  in 

dorsal  view  Triepicharis 

7a.  Dorsal  face  of  scutellum  flat;  prepygidial  fimbria  of  fe- 
male preceded  by  a shiny,  nearly  glabrous  area  ...  8 
b.  Dorsal  face  of  scutellum  bigibbous;  female  without  shiny, 

glabrous  area  anterior  to  prepygidial  fimbria 

Cyphepicharis 

8a.  Jugal  lobe  of  posterior  wing  about  half  as  long  as  vannal 
lobe  and  nearly  attaining  apex  of  cubital  cell;  female 
metabasitarsus  about  twice  longer  than  broad,  posterior 
margin  nearly  straight;  male  pygidial  plate  broad,  cov- 
ering most  of  seventh  tergum  Epicharoides 

b.  Jugal  lobe  of  posterior  wing  about  one-third  as  long  as 
vannal  lobe,  its  apex  well  short  of  that  of  cubital  cell; 
female  metabasitarsus  about  one-third  longer  than  broad, 
posterior  margin  strongly  curved;  male  pygidial  plate 
narrow,  covering  less  than  one-half  of  seventh  tergum 
Epicharitides 

Subgenus  Epicharana  Michener 

Epicharis:  Moure,  1 945a:294-295.  Type-species:  " Epicharis 
rustica  Olivier,  1 789”  = Apis  rustica  Olivier,  1789.  Not 
Epicharis  Klug,  1 807. 

Epicharis  subg.  Epicharana  Michener,  1954:144.  Type- 
species:  Apis  rustica  Olivier,  1789;  original  designation. 

DESCRIPTION 

Maxillary  palp  three-segmented,  third  segment  a little  short- 
er, and  much  narrower,  than  second;  lateral  ridge  of  clypeal 
disc  strong;  malar  area  about  as  long  as  minimum  thickness 
of  first  flagellar  segment;  occipital  margin  rounded;  occipital 
fiagelliform  setae  long,  extending  beyond  anterior  margin  of 
tegula;  metanotum  bifaced,  dorsal  face  about  as  long  as  pos- 
terior face,  juncture  angular  or  crested;  jugal  lobe  of  posterior 
wing  shorter  than  cubital  cell  and  less  than  half  as  long  as 
vannal  lobe. 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  51 


Female.  Labrum  with  low,  median  longitudinal  ridge;  out- 
er face  of  mesobasitarsus,  on  anterior  one-third  with  mixed 
long,  coarse,  simple  setae  and  shorter,  fine,  plumose  hairs, 
posterior  two-thirds  with  sparse  longer,  coarse,  simple  setae 
and  sparse,  short,  fine,  plumose  hairs;  basitibial  plate  with 
secondary  plate;  metatibia  no  longer  than  metabasitarsus; 
third  and  fourth  terga  without  basal  specialized  areas;  in 
dorsal  view,  margins  of  pygidial  plate  concave,  apex  broadly 
truncate. 

Male.  Labrum  without  median  ridge;  first  flagellar  segment 
shorter  than  scape,  longer  than  second  segment,  much  shorter 
then  second  and  third  combined;  ocellocular  distance  greater 
than  ocellar  diameter;  procoxa  with  flattened  distal  process; 
mesosternum  with  prominent  process  on  each  side  of  mid- 
line,  anterior  to  mesocoxa;  metatrochanter  and  base  of  meta- 
femur with  ventral  mat  of  short  blackish  setae;  metatibia 
with  carinate  posteroventral  ridge;  metabasitarsus  carinate 
along  anterior  margin,  ending  in  prominent  tooth-like  pro- 
cess at  apex  of  segment;  pygidial  plate  broad,  its  margins 
confluent  with  those  of  segment,  apex  bidentate. 

This  is  the  group  that  Moure  (1945a)  incorrectly  inter- 
preted as  Epicharis  s.s.,  as  discussed  above.  Five  species 
occur  in  Central  America,  with  a few  more  in  South  America. 


KEY  TO  CENTRAL  AMERICAN  EPICHARANA 

la.  Abdominal  terga  blackish  and  immaculate  (except  male 
of  one  species  with  dorsal  fascia  on  first  and  lateral  spot 
on  second)  or  ferruginous  and  immaculate;  male  clypeus 

black 2 

b.  Abdominal  terga  reddish  brown  to  ferruginous  and  dor- 
sal face  of  first  segment  with  narrowly  interrupted  trans- 
verse yellow  fascia  (sometimes  obscure);  male  clypeus 

yellow elegans  F.  Smith 

2a.  Dorsal  segments  of  abdomen  black,  first  tergite  with  or 
without  transverse  yellow  fascia  on  dorsum;  male  meta- 
tibia moderately  convex  in  middle  of  anterior  margin 
(Fig.  66);  disc  of  female  clypeus,  in  basal  one-third  or 
more,  with  more  or  less  distinct,  weakly  depressed,  me- 
dian impunctate  line  3 

b.  Dorsal  segments  of  abdomen  bright  ferruginous  and  im- 
maculate; male  metatibia  strongly,  obtusely  convex  in 
middle  of  anterior  margin  (Fig.  65);  disc  of  female  clyp- 
eus uniformly,  subcontiguously  punctate  in  basal  one- 

third  bova,  new  species 

3a.  Female:  scutellum  shinier  than  mesoscutum,  punctures 
of  disc  both  less  distinct  and  more  separated  than  those 
of  mesoscutum;  larger  punctures  of  parapsis  sparse,  sep- 
arated by  more  than  twice  a puncture  diameter.  Male: 
first  tergum  without  dorsal  transverse  yellow  fascia,  me- 
sosternal  tubercles  nearly  prostrate,  their  inner  margins 

rounded  (Fig.  67) rustica  (Olivier) 

b.  Female:  scutellum  no  shinier  than  mesoscutum,  micro- 
punctures of  disc  as  sharp  and  dense  as  those  of  meso- 
scutum; larger  punctures  of  parapsis  dense,  separated  by 
about  a puncture  diameter  or  less.  Male:  first  tergite  with 
narrowly  interrupted  yellow  fascia  on  dorsal  face;  me- 


sostemal  tubercles  erect,  obtuse,  inner  margin  sharply 
carinate  (Fig.  68)  angulosa,  new  species 

Epicharis  ( Epicharana ) angulosa,  new  species 

Figures  66,  68,  69-72 

DIAGNOSIS 

Separable  from  other  species  of  Epicharana  by  the  following 
unique  combination  of  features:  pubescence,  except  of  hind 
legs,  black;  abdomen  black,  except  dorsally  on  first  tergum 
and  laterally  on  second  tergum  of  male.  Male:  mesosternal 
tubercles  angulate  along  inner  margin,  high,  long  and  obtuse 
in  profile;  metatibia  moderately  swollen  (Fig.  66),  posterior 
ventral  carina  low,  abruptly  reduced  beyond  midlength.  Fe- 
male: shiny  area  at  base  of  clypeal  disc  no  greater  in  area 
than  triangular  supraclypeal  area  and  basal  one-half  of  disc 
densely  and  coarsely  punctate,  without  definite  impunctate 
median  line. 

DESCRIPTION 

HOLOTYPE  MALE.  Measurements  (mm).  Head  width 
5.95  (5.64-5.90);  head  length  4.00  (3.90-4.00);  wing  length 
17.0  (15.0-16.5);  total  length  21.0  (20.0-22.0). 

Head.  1 .48  ( 1 .45-1.49)  times  broader  than  long;  in  frontal 
view,  occipital  margin  nearly  straight  (except  ocellar  eleva- 
tion) and  slightly  below  level  of  tops  of  eyes;  inner  eye  mar- 
gins moderately  convergent  above,  upper  frontal  width  0.77 
(0.76-0.79)  times  lower  frontal  width.  Mandible  slender  be- 
yond middle,  inner  tooth  large,  blunt.  Labrum  quadrate, 
slightly  broader  than  long,  apical  margin  transverse,  disc  shiny 
and  weakly  tessellate  between  sparse  to  close  punctures  vary- 
ing from  fine  to  coarse.  Clypeus  about  1 .7  times  broader  than 
long,  apicolateral  angle  removed  from  eye  by  slightly  less 
than  diameter  of  antennal  socket;  disc  moderately  shiny  be- 
tween dense,  coarse  punctures  except  near  apical  margin  where 
punctures  are  sparse  and  fine  and  integument  is  more  or  less 
“wrinkled,”  median  impunctate  line  absent;  discal  carinae 
moderately  convergent  above,  distance  between  them  at  low- 
er end  about  1.7  times  that  at  upper  end;  side  slightly  shiny 
and  distinctly  roughened  between  variably  sparse  to  subcon- 
tiguous,  fine  to  coarse  punctures.  Frons  moderately  shiny 
and  distinctly  tessellate  between  dense,  mixed  fine  and  mod- 
erate punctures;  preocellar  area  slightly  bulging  on  either  side, 
smooth,  shiny,  and  impunctate;  ocellocular  area  dull  and 
densely  tessellate,  subcontiguously  micropunctate  and  with 
scattered  minute  punctures;  preocciput  moderately  shiny  be- 
tween dense,  fine  punctures;  gena  moderately  shiny  to  shiny 
between  close  to  dense  punctures,  minute  near  eye,  becoming 
fine  near  margin.  Interantennal  distance  1.86  (1.88-2.05) 
times  antennal  socket  diameter;  antennocular  distance  0.59 
(0.63-0.71)  times  antennal  socket  diameter;  scape  robust, 
1.50  (1.50-1.64)  times  longer  than  broad,  scape  length  1.57 
(1.53-1.61)  times  length  of  first  flagellar  segment;  first  fla- 
gellar segment  1.31  (1 .22-1 .32)  times  longer  than  broad,  much 
shorter  than  following  three  segments  combined,  1.15(1 .08- 
1.13)  times  longer  than  second.  Interocellar  distance  1.70 
(1.44-1.64)  times  diameter  of  anterior  ocellus;  ocellocular 


52  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


Figures  63-64,  pygidial  plate  of  female:  63,  Epicharis  ( Epicharana ) rustica\  64,  E.  ( Hoplepicharis ) lunulata.  Figs.  65-66,  metatibia  of  male: 
65,  E.  ( Epicharana ) bova\  66,  E.  (Epicharana)  angulosa.  Figs.  67-68,  profile  of  right  mesosternal  process  of  male:  67,  E.  ( Epicharana ) rustica\ 
68,  E.  (Epicharana)  angulosa. 


distance  1 .96  ( 1 .89-1 .93)  times  diameter  of  anterior  ocellus; 
ocelloccipital  distance  1.70  (1.50-1.67)  times  diameter  of 
anterior  ocellus. 

Thorax.  Mesoscutum  slightly  shiny,  uniformly  densely, 
finely  punctate  and  with  sparse  moderate  punctures  which 
become  scattered  distad;  dorsal  face  of  scutellum  broadly 
depressed  in  middle,  punctation  as  scutum,  but  fine  punc- 
tures scattered;  metanotum  moderately  shiny  in  median  area, 
dull  elsewhere,  roughened  and  tessellate,  with  minute  obscure 
punctures.  Mesepisternum  moderately  shiny  and  tessellate 
between  sparse,  shallow,  fine  punctures;  metepisternum 
moderately  shiny  between  subcontiguous,  minute  punctures 


(appearing  finely  reticulopunctate  at  certain  angles).  Meso- 
sternal process,  in  profile,  suberect  and  obtusely  triangular 
(Fig.  68);  in  ventral  view,  inner  margin  carinate.  Propodeum 
moderately  shiny,  minutely  reticulopunctate  and  with  sparse 
to  scattered  fine  punctures.  Procoxal  process  broadly  ellip- 
tical, with  acute  apex;  mesofemur  stout,  twice  longer  than 
thick,  greatest  thickness  a little  basad  of  midlength;  antero- 
basal  ventral  depression  of  metafemur  subcarinate  along  its 
posterior  margin;  anterior  edge  of  metatibia  evenly  curved 
(Fig.  66),  posterior  carina  low  throughout,  abruptly  reduced 
distad  of  middle. 

Abdomen.  Tergal  discs  moderately  shiny  between  minute 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  53 


sSSwrosj 


Figures  69-72.  Epicharis  ( Epicharana ) angulosa,  male  seventh  and  eight  stermtes  and  genitalia  (ventral  and  dorsal  views).  Scale  line  = 1.00 
m m . 


54  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


punctures  which  are  dense  on  basal  segments,  becoming  pro- 
gressively coarser  and  less  close  on  succeeding  segments;  discs 
also  with  scattered  fine  punctures  on  basal  segments,  becom- 
ing progressively  coarser  (but  still  fine)  on  succeeding  seg- 
ments. Seventh  tergite  weakly  raised  in  middle  (broader  ba- 
sad)  to  form  a weakly  differentiated  median  plate;  apex 
bidentate,  teeth  long,  slender,  acute,  emargination  longer  than 
a semicircle. 

Terminalia.  Apical  margin  of  distal  lobe  of  seventh  sternite 
(Fig.  69)  broadly,  shallowly  concave;  setae  long,  some  con- 
spicuously plumose.  Shoulders  of  disc  of  eighth  sternite  (Fig. 
70)  angular  and  sides  of  apical  lobe  of  disc  strongly  conver- 
gent. Genitalia  as  in  Figs.  71  and  72. 

Pilosity.  Generally  blackish  brown;  sides  of  pronotal  collar, 
pronotal  lobe,  mesoscutum  (especially  anteriorly  and  at  side) 
with  admixed  very  pale  brown  hairs;  hairs  on  side  of  tergites 
and  on  abdominal  venter  mixed  medium  and  light  brown. 
Flairs  mostly  long,  erect,  plumose  and  dense,  but  labrum 
nearly  bare.  Metatibia  and  metabasitarsus  with  long  yellow- 
ish hairs.  Abdominal  terga,  beyond  first,  with  sparse  discal 
hairs  which  are  short,  simple  and  suberect  on  second  seg- 
ment, becoming  progressively  longer  and  more  abundant 
caudad;  terga  also  with  abundant  minute,  appressed,  scale- 
like hairs  (arising  from  minute  punctures)  which  become 
progressively  longer  on  succeeding  segments. 

Color.  Blackish  brown.  The  following  pale  yellow:  oblique, 
linear  mark  near  base  of  mandible;  labrum;  oblique,  linear 
mark  from  clypeal  margin,  at  tentorial  pit,  to  malar  area; 
triangular  supraclypeal  mark;  underside  of  scape;  spot  on 
procoxal  process;  anterodistal  spot  on  pro-  and  mesofemora; 
large  distal  spot  on  metafemur;  stripe  on  anterior  margin  of 
protibia  nearly  to  apex;  broad  stripe  on  anterior  margin  of 
mesotibia  nearly  to  apex,  constricted  in  middle;  anterior  and 
outer  face  of  metabasitarsus;  large,  anterior  spots  on  dorsal 
face  of  first  tergum,  narrowly  separated  in  middle;  small  lat- 
erobasal  spot  on  second  tergum.  Underside  of  flagellum  light 
brown;  apical  tarsal  segments  reddish.  Wings  uniformly  dark 
brown,  veins  and  stigma  blackish. 

FEMALE.  Measurements  (mm).  Head  width  6.31;  head 
length  4.15;  wing  length  16.5;  total  length  23.0. 

Head.  1.52  times  broader  than  long;  in  frontal  view,  oc- 
cipital margin  straight,  except  ocellar  elevation;  inner  eye 
margins  moderately  convergent  above,  upper  frontal  width 
0.84  times  lower  frontal  width.  Mandible  stout,  tridentate, 
inner  teeth  obtuse  and  margin  between  them  broadly  con- 
cave. Labrum  about  1 . 1 times  longer  than  broad,  apex  sub- 
angularly  rounded  in  middle;  disc  moderately  shiny,  rough- 
ened and  tessellate  between  dense  to  subcontiguous,  mixed 
fine  and  coarse  punctures,  median  line  slightly  raised.  Clyp- 
eus  about  1.6  times  broader  than  long,  apicolateral  angle 
separated  from  eye  by  0.5  times  diameter  of  antennal  socket; 
discal  carinae  moderately  convergent  basad,  intercarinal  dis- 
tance at  base  about  0.66  times  that  at  their  distal  end;  sculp- 
ture as  described  for  male,  but  with  shiny  basal  area  which 
is  no  greater  than  supraclypeal  area.  Punctation  of  frons, 
preoccipital  area,  and  gena  as  in  male.  Interantennal  distance 
2.20  times  antennal  socket  diameter;  antennocular  distance 
0.88  times  antennal  socket  diameter;  scape  robust,  1 .9 1 times 


longer  than  wide,  scape  length  1.05  times  length  of  first  fla- 
gellar segment;  first  flagellar  segment  shorter  than  following 
three  combined,  3.81  times  longer  than  second.  Interocellar 
distance  1 .44  times  diameter  of  anterior  ocellus;  ocellocular 
distance  1.78  times  diameter  of  anterior  ocellus;  ocelloccip- 
ital  distance  1.56  times  diameter  of  anterior  ocellus. 

Thorax.  Punctation  as  in  male.  Metanotum  with  horizon- 
tal basal  face  separated  from  vertical  posterior  face  by  a 
cariniform  ridge  on  either  side  of  middle.  Apex  of  basitibial 
plate  acute;  secondary  plate  about  twice  longer  than  wide. 

Abdomen.  Punctation  as  described  for  male.  Disc  of  py- 
gidial  plate  not  visible. 

Pilosity.  About  as  described  for  male,  but  pronotal  collar 
and  lobe  without  pale  hairs;  bristles  of  thoracic  venter  with 
pale  tips;  scale-like  hairs  of  fifth  tergum  becoming  longer, 
more  erect  and  plumose  toward  prepygidial  fringe;  hairs  of 
prepygidial  fringe  reddish  brown  and  pale  tips;  scopa!  hairs 
pale  yellowish. 

Color.  Generally  blackish  brown;  mandible  (except  golden 
apical  blotch),  underside  of  flagellum,  apical  segments,  red- 
dish brown;  paraocular  area  with  small,  obscure  yellowish 
blotch  near  clypeal  margin,  below  level  of  tentorial  pit  (larger 
and  more  distinct  on  left  side).  Wings  as  in  male. 

TYPE  MATERIAL 

Holotype  male,  allotype,  and  one  male  paratype:  Monte- 
verde,  1500  m elev.,  Puntarenas  Prov.,  COSTA  RICA,  12 
Sept.  1978  (G.W.  Frankie),  in  LACM.  Paratypes:  13,  same 
locality,  19  Aug.  1974  (D.  Janzen;  UKAN);  233,  4 km  E San 
Ignacio  de  Acosta,  4000  ft.  elev.,  San  Jose  Prov.,  COSTA 
RICA,  8 July  1963  (C.D.  Michener  et  al.;  UKAN). 

ETYMOLOGY 

The  specific  epithet  is  a Latin  word  meaning  with  corners, 
referring  to  the  angulate  mesosternal  processes  of  the  male. 

DISCUSSION 

Aside  from  the  variations  noted  above  in  the  measurements 
and  proportions,  the  males  available  are  very  similar  to  one 
another.  The  laterobasal  spot  on  the  second  tergite  may  ex- 
tend mesad  as  a narrow  line  and/or  a short  distance  distad. 
The  procoxal  process  is  immaculate  in  three  paratypes  and 
in  these  specimens  the  mesotibial  stripe  is  divided  into  a 
basal  spot  and  two  narrowly  separated  stripes  along  the  seg- 
ment. One  male  has  a narrow  yellow  stripe  on  the  anterior 
face  of  the  metadistitarsus  and  yellow  blotches  on  the  outer 
face  of  the  metamediotarsi. 

Males  of  this  species  are  easily  recognized  by  the  combi- 
nation of  black  abdomen  with  contrasting  pale  marks,  sub- 
erect and  internally  carinate  mesosternal  tubercles  and  black 
labrum.  Females  are  considerably  less  distinctive,  but  differ 
from  such  species  as  E.  elegans,  E.  rustica,  and  E.  bova  in 
possessing  an  immaculate  abdomen  and  largely  black  tho- 
racic pubescence.  The  lack  of  a median  impunctate  line  on 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  55 


Figures  73-76.  Epicharis  (Epicharana)  bova,  male  seventh  and  eighth  sternites  and  genitalia  (ventral  and  dorsal  views).  Scale  line  = 1.00  mm. 


56  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


the  clypeal  disc  will  also  separate  E.  angulosa  from  E.  elegans 
and  E.  rustica. 

Epicharis  ( Epicharana ) bova,  new  species 

Figures  65,  73-76 

DIAGNOSIS 

Separable  from  all  other  species  of  Epicharana  by  the  fol- 
lowing unique  combination  of  characters.  Abdomen  ferru- 
ginous, immaculate;  pubescence  of  head  and  thorax  medium 
brown  to  blackish  brown,  mesoscutum  without  pale  hairs; 
clypeus  black.  Male:  mesostemal  processes,  in  profile,  sub- 
erect but  low  and  rounded,  not  carinate  on  inner  margin; 
anterior  margin  of  metatibia  strongly  produced  (Fig.  65). 
Female:  clypeal  disc  subcontiguously  to  densely  punctate, 
without  median  impunctate  line. 

DESCRIPTION 

1 IOLOTYPE  MALE.  Measurements  (mm).  Head  width 
5.95;  head  length  4.15;  wing  length  16.0;  total  length  25.0. 

Head.  1.43  times  broader  than  long;  in  frontal  view,  oc- 
cipital margin  very  weakly  concave,  except  for  ocellar  ele- 
vation; ocelli  on  occipital  margin;  upper  frontal  width  0.71 
times  lower  frontal  width.  Mandible  slender,  inner  tooth  small, 
obtuse.  Labrum  about  1.2  times  broader  than  long,  apical 
margin  broadly  rounded;  disc  moderately  shiny,  weakly  tes- 
sellate  between  irregularly  spaced  (mostly  sparse),  fine  and 
coarse  punctures.  Clypeus  about  1 . 5 times  broader  than  long, 
apicolateral  angle  separated  from  inner  eye  margin  by  about 
0.8  times  diameter  of  antennal  socket;  clypeus  shiny  and  very 
weakly  tessellate  between  subcontiguous  to  dense,  coarse 
punctures  which  become  moderate  distad,  median  impunc- 
tate line  absent;  discal  carinae  about  twice  as  far  apart  distad 
as  at  base;  side  shiny  between  dense,  moderate  to  coarse, 
elongate  (especially  distad)  punctures.  Frons  moderately  shiny 
and  distinctly  tessellate  between  dense,  mixed  fine  and  mod- 
erate punctures;  preocellar  area  slightly  protuberant  on  either 
side,  shiny  and  nearly  impunctate;  ocellocular  area  dull  and 
densely  tessellate,  subcontiguously,  shallowly  micropunctate 
and  with  scattered  minute  punctures;  preocciput  moderately 
shiny  to  shiny  between  close  to  dense  punctures,  minute  near 
eye,  becoming  fine  near  margin.  Interantennal  distance  2.00 
times  antennal  socket  diameter;  antennocular  distance  0.63 
times  antennal  socket  diameter;  scape  robust,  1.74  times 
longer  than  wide,  scape  length  1.52  times  length  of  first  fla- 
gellar segment;  first  flagellar  segment  shorter  than  following 
three  segments  combined,  1.37  times  longer  than  second 
segment.  Interocellar  distance  1.61  times  diameter  of  anterior 
ocellus;  ocellocular  distance  1.96  times  diameter  of  anterior 
ocellus;  ocelloccipital  distance  1.61  times  diameter  of  ante- 
rior ocellus. 

Thorax.  Mesoscutum  slightly  shiny,  uniformly  densely, 
finely  punctate,  and  with  sparse,  moderate  punctures  which 
become  scattered  distad;  dorsal  face  of  scuteilum  broadly 
depressed  in  middle,  punctures  as  on  mesoscutum,  but  fine 
punctures  scattered;  dorsal  face  of  metanotum  moderately 
shiny  and  distinctly  tessellate  between  dense,  fine  punctures. 


posterior  face  dull,  densely  tessellate  and  with  scattered  fine 
punctures.  Mesepisternum  moderately  shiny  and  tessellate 
between  sparse,  fine  punctures;  metepisternum  moderately 
shiny  between  subcontiguous,  minute  punctures.  Mesoster- 
nal  process  suberect  in  profile,  its  apex  narrowly  rounded;  in 
ventral  view,  inner  margin  rounded.  Propodeum  moderately 
shiny,  subcontiguously  micropunctate  between  sparse  to 
scattered,  fine  punctures.  Procoxal  process  subcircular,  apex 
not  produced;  mesofemur  stout,  about  twice  longer  than  thick, 
greatest  thickness  a little  basad  of  midlength;  anterobasal 
ventral  depression  of  metafemur  subcarinate  along  its  pos- 
terior margin;  anterior  margin  of  metatibia  abruptly  convex 
at  about  midlength  and  anterior  carina  plainly  visible  (Fig. 
65),  posterior  carina  low  throughout,  highest  distad  of  middle 
and  abruptly  reduced  beyond  highest  point. 

Abdomen.  Tergal  discs  moderately  shiny  between  minute 
punctures  which  are  dense  on  basal  segments,  becoming  pro- 
gressively coarser  and  less  close  on  succeeding  segments;  discs 
also  with  scattered  fine  punctures  on  basal  segments,  becom- 
ing progressively  coarser  (but  still  fine)  on  succeeding  seg- 
ments. Seventh  tergite  with  very  weakly  differentiated  py- 
gidial  plate  which  is  depressed  along  midline;  apex  bidentate, 
teeth  stout  and  acute,  emargination  between  them  semicir- 
cular. 

Terminalia.  Apical  margin  of  distal  lobe  of  seventh  sternite 
(Fig.  73)  deeply,  angularly  incised;  setae  long,  widely  spaced, 
mostly  simple.  Shoulders  of  disc  of  eighth  sternite  (Fig.  74) 
obtuse  and  sides  of  apical  lobe  of  disc  weakly  convergent 
distad.  Genitalia  as  in  Figs.  75  and  76. 

Pilosity.  Generally  blackish  brown  on  head,  thorax,  and 
legs,  but  yellowish  on  metatibia  and  metabasitarsus;  suberect 
hairs  and  appressed  scaliform  hairs  of  terga  two  to  six  golden 
brown,  but  fringes  of  fifth  and  sixth  segments  brown  and  a 
few  brown,  simple,  suberect  hairs  scattered  on  discs.  Scale- 
like  hairs  very  short  basad,  becoming  longer  and  more  erect 
on  succeeding  segments.  Hairs  of  abdominal  sterna  golden 
brown  to  blackish  brown. 

Color.  Head,  thorax,  and  legs  blackish  brown;  abdominal 
terga  ferruginous,  sterna  light  brown.  The  following  yellow- 
ish: large  triangular  mark  near  base  of  mandible;  labrum; 
narrow,  oblique  stripe  on  side  of  face,  from  tentorial  pit  to 
malar  area;  small,  triangular  supraclypeal  spot;  small  spot  on 
underside  of  scape;  basal  spot  on  protibia;  dorsal,  apical  spot 
on  metafemur;  outer  face  of  metatibia  and  metabasitarsus 
(metatibia  black  at  base).  Inner  leg  surfaces  dark  ferruginous; 
tarsi  ferruginous.  Wings  blackish  brown,  veins  and  stigma 
black. 

FEMALE.  Measurements  (mm).  Head  width  6.41;  head 
length  4.26-4.31;  wing  length  16.5;  total  length  24.0-27.0. 

Head.  1.49-1.51  times  broader  than  long;  in  frontal  view, 
occipital  margin  nearly  straight,  ocellar  elevation  anterior  to 
margin;  upper  frontal  width  0.79-0.81  times  lower  frontal 
width.  Mandible  stout,  tridentate,  inner  teeth  obtuse,  margin 
between  them  broadly  concave.  Labrum  about  1.1  times 
longer  than  broad,  apex  subangularly  rounded  in  middle;  disc 
moderately  shiny,  roughened,  and  tessellate  between  dense 
to  subcontiguous,  mixed  fine  and  coarse  punctures,  median 
line  slightly  raised.  Clypeus  about  1.4  times  broader  than 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  57 


long,  apicolateral  angle  separated  from  inner  eye  margin  by 
about  0.75  times  antennal  socket  diameter;  intercarinal  dis- 
tance at  base  about  0.36  times  that  at  their  distal  end;  sculp- 
ture as  described  for  male.  Remaining  cephalic  sculpture  as 
described  for  male.  Interantennal  distance  2.28-2.37  times 
antennal  socket  diameter;  antennocular  distance  0.92-0.95 
times  antennal  socket  diameter;  scape  robust,  1 .86-1.9 1 times 
longer  than  broad,  scape  length  1.04-1.08  times  length  of 
first  flagellar  segment;  first  flagellar  segment  shorter  than  fol- 
lowing three  segments  combined,  3.45-3.85  times  length  of 
second  segment. 

Thorax.  Punctation  as  in  male.  Dorsal  and  posterior  faces 
of  metanotum  separated  on  either  side  of  middle  by  short, 
convex,  cariniform  ridge.  Basitibial  plate  subacute  at  apex, 
secondary  plate  about  2.8  times  longer  than  wide. 

Abdomen.  Punctation  as  in  male.  Pygidial  plate  broadly 
truncate  at  apex,  secondary  plate  very  weak. 

Pilosity.  About  as  described  for  male,  with  following  dif- 
ferences: discs  of  third,  fourth,  and  fifth  terga  with  some 
suberect  to  erect  dark  brown,  bristle-like  hairs,  longer  and 
more  abundant  on  succeeding  segments;  prepygidial  fringe 
golden  brown.  Scopa  of  metatibia  and  metabasitarsus  yel- 
lowish. 

Color.  As  described  for  male,  but  wholly  without  yellowish 
markings  and  pronotum  and  propodeum  reddish  brown. 

TYPE  MATERIAL 

Holotype  male:  Cerro  Campana,  Panama  Prov.,  PANAMA, 
4 May  1960  (W.J.  Hanson),  in  Snow  Entomological  Museum, 
University  of  Kansas.  Allotype:  4 mi.  S San  Vito  de  Java, 
Puntarenas  Prov.,  COSTA  RICA,  15  Aug.  1967  (R.W. 
McDiarmid;  LACM).  Paratypes:  IS,  N of  El  Valle  de  Anton, 
Code  Prov.,  PANAMA,  12-13  Sept.  1964  (R.L.  Dressier; 
UKAN);  IS,  O.T.S.  Field  Station,  Finca  La  Selva,  Heredia 
Prov.,  COSTA  RICA,  mid  Aug.  1980  (J.M.  MacDougal; 
LACM),  on  Passiflora  lobata,  0900. 

ETYMOLOGY 

The  specific  epithet  is  a Latin  word  for  a swelling  of  the  legs 
and  alludes  to  the  swollen  metatibia  of  the  male. 

DISCUSSION 

Both  sexes  are  easily  separated  from  others  in  the  subgenus 
Epicharana  by  the  features  listed  above  in  the  Diagnosis. 
The  male  is  especially  distinctive  in  the  shape  of  the  meta- 
tibia, the  anterior  margin  of  which  is  conspicuously  and 
abruptly  convex  at  about  midlength.  The  low  posterior  carina 
of  the  metatibia  is  like  that  of  E.  angulosa,  as  is  the  poste- 
riorly subcarinate  anterobasal  depression  on  the  underside 
of  the  metafemur.  The  abdomen  of  E.  angulosa  is  black  and 
the  mesosternal  tubercles  are  acute  at  their  apices  and  sharply 
carinate  on  their  inner  margins. 

The  only  other  species  with  a reddish  abdomen  is  E.  ele- 
gans,  in  both  sexes  of  which  the  abdomen  is  distinctly  yel- 
lowish red  and  the  first  tergum  is  yellowish  maculate  on  the 
disc.  In  males  of  E.  elegans,  and  those  of  E.  flava  and  E. 


rustica,  the  clypeus  is  pale,  the  metatibia  is  regularly  convex 
along  its  anterior  margin,  there  is  no  inner,  anterior  meta- 
tibial  carina,  and  the  inner  posterior  metatibial  carina  is  ex- 
panded beyond  midlength  and  folded  mesad.  In  both  sexes 
of  these  species,  E.  elegans,  E.  Jlava,  and  E.  rustica,  there  is 
considerable  pale  pubescence  intermixed  with  blackish  on 
the  thoracic  dorsum.  Females  of  these  three  species  have  a 
definite  median  impunctate  line  on  the  clypeus  and  the  mi- 
nute punctures  of  the  scutellum  are  usually  coarser  and  less 
distinct  than  those  of  the  mesoscutum. 

Epicharis  ( Epicharana ) elegans  F.  Smith 

Epicharis  elegans  F.  Smith,  1 86 1 : 1 52.  9 <5. 

Epicharis  elegans:  Moure,  1945a:296  (tax.). 

Epicharis  salazari  Cockerell,  1949:480-481.  9.  NEW  SYN- 
ONYMY. 

This  is  a common  black  and  red  species  in  Mexico  and  ranges 
south  to  Costa  Rica.  Specimens  from  southern  Mexico  (Chia- 
pas and  Yucatan),  Guatemala,  El  Salvador,  and  Costa  Rica 
have  the  abdomen  more  brownish  and  correspond  to  E. 
salazari,  described  from  El  Salvador.  These  are  minor  vari- 
ants and  not  worthy  of  separation  from  E.  elegans. 

NEW  RECORDS 

MEXICO,  CHIAPAS:  499,  La  Revancha,  20  Aug.  1972  (T.W. 
Taylor;  LACM).  GUERRERO:  699,  12.7  mi.  N Iguala,  5200 
ft.  elev.,  1 Aug.  1969  (Univ.  Kans.  Mex.  Exped.;  UKAN); 
19,  23  mi.  N Taxco,  1 700  ft.  elev.,  8 Aug.  1 954  (Univ.  Kans. 
Mex.  Exped.;  UKAN);  19,  3.7  mi.  W Rio  Balsas,  5 Aug.  1 965 
(G.W.  Byers  and  party;  UKAN);  2 33,  Iguala,  2400  ft.  elev., 
8 Aug.  1954  (Univ.  Kans.  Mex.  Exped.;  UKAN).  JALISCO: 

18,  Guadalajara,  no  date  (Crawford;  LACM);  399,  Cocula, 
4450  ft.  elev.,  27  Sept.  1957  (H.A.  Scullen;  ORSU);  299, 
Puente  Grande,  5000  ft.  elev.,  20  Aug.  1954  (Univ.  Kans. 
Mex.  Exped.;  UKAN).  MORELOS:  599,  888,  Lake  Tequis- 
quetengo,  5000  ft.  elev.,  13  Sept.  1957  (H.A.  Scullen;  ORSU); 
13,  Rancho  Tetela,  Cuernavaca,  24  June  1974  (K.E.  Donahue 
and  S.  Adams;  LACM);  19,  13,  Cuernavaca,  no  date  (Craw- 
ford; LACM);  233,  Alpuyeca,  27  June  and  3 July  1951  (P.D. 
Hurd;  UCB);  19,  7.3  mi.  S Yautepec,  3000  ft.  elev.,  16  Aug. 
1962  (Ordway  and  Naumann;  UKAN);  399,  4 mi.  SW  Yau- 
tepec, 3800  ft.  elev.,  2 July  1961  (C.D.  Michener;  UKAN), 
on  Cassia  sp.;  299,  4.3  mi.  W Yautepec,  4000  ft.  elev.,  17 
Aug.  1962  (Ordway  and  Marston;  UKAN);  13,  7 mi.  NE 
Yautepec,  4000  ft.  elev.,  18  Aug.  1962  (Univ.  Kans.  Mex. 
Exped.;  UKAN);  13,  14  mi.  S Yautepec,  16  Aug.  1962  (Mar- 
ston and  Roberts;  UKAN).  OAXACA:  19,  25  mi.  SE  Oaxaca, 
5600  ft.  elev.,  27  June  1963  (Scullen  and  Bolinger;  ORSU); 

19,  13,  47  mi.  SE  Oaxaca,  13  July  1952  (E.E.  Gilbert  and 
C.D.  MacNeil;  UCB);  2 99,  5 mi.  NW  Totolapan,  4000  ft. 
elev.,  29  July  1970  (E.M.  Fisher  and  P.  Sullivan;  LACM); 
4799,  5 mi.  NW  Totolapan,  3800  ft.  elev.,  6 July  1 953  (Univ. 
Kans.  Mex.  Exped.;  UKAN),  on  Malpighia  mexicana ; 19, 
Salina  Cruz,  no  date  (F.K.  Knab;  USNM);  299,  Mixtla,  5600 
ft.  elev.,  22  Aug.  1963  (Scullen  and  Bolonger;  ORSU);  433, 
Monte  Alban,  6000  ft.  elev.,  27  June  196 1 (Univ.  Kans.  Mex. 


58  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


Exped.;  UKAN);  1 <3,  same,  except  15  July  1955;  19,  Tama- 
zulapan,  6000  ft.  elev.,  28  June  1961  (Univ.  Kans.  Mex. 
Exped.;  UKAN);  19,  2 mi.  NW  Tamazulapan,  6000  ft.  elev., 
28  June  1961  (Univ.  Kans.  Mex.  Exped.;  UKAN).  PUEBLA: 
999,  22  km  NW  Izucar  de  Matamoros,  1158m  elev.,  2 1 Sept. 
1976  (C.D.  George  and  R.R.  Snelling;  LACM),  on  Cassia 
laevigata ; 1699,  16.1  km  NW  Izucar  de  Metamoros,  1280  m 
elev.,  1 7 Sept.  1976  (C.D.  George  and  R.R.  Snelling;  LACM), 
on  Caesalpinia  cacalaco\  999,  788,  6.9  km  S Izucar  de  Mat- 
amoros, 1 250  m elev.,  1 7 Sept.  1 976  (C.D.  George  and  R.R. 
Snelling;  LACM),  on  Solatium  rostratum  (99)  and  Martynia 
annua  (88);  399,  388,  Atlixco,  9 July  1970  (R.E.  Beer  and 
party;  L1KAN);  299,  3 mi.  NW  Petlalcingo,  4600  ft.  elev.,  29 
Aug.  and  5 Sept.  1 972  (Byers  and  Thornhill;  UKAN);  19,  12 
mi.  NW  Tehuitzingo,  4050  ft.  elev.,  29  June  1961  (Univ. 
Kans.  Mex.  Exped.;  UKAN);  on  Cassia  sp.;  19,  10  mi.  SE 
Tehuitzingo,  3900  ft.  elev.,  3 July  1953  (Univ.  Kans.  Mex. 
Exped.;  UKAN).  SAN  LUIS  POTOSI:  688,  El  Salto,  1600 
ft.  elev.,  24  Aug.  1954  (Univ.  Kans.  Mex.  Exped.;  UKAN). 
TAMAULIPAS:  19,  38  mi.  N El  Mante,  1050  ft.  elev.,  1 1 
Oct.  1957  (H.A.  Scullen;  ORSU).  YUCATAN:  599,  Piste,  24 
June  1967  (E.C.  Welling;  LACM).  GUATEMALA:  299,  “1923 
F.4696”  (UKAN);  19,  “env.  de  Guatemala”  (R.  Guerin; 
MNHN).  EL  SALVADOR:  19,  Dept.  Santa  Tecla,  Feb.  1947 
(M.  Salazar;  USNM;  type  of  E.  salazari)]  499,  Santa  Tecla, 
900  m elev.,  25  Apr. -6  May  1972  (S.  and  L.  Steinhaus; 
DPIF).  COSTA  RICA,  GUANACASTE:  19,  Comelco,  8 km 
NW  Bagaces,  3 1 March  1971  (P.S.  Opler;  UCB),  on  Tabebuia 
rosea\  19,  same,  except  25  Nov.  1972;  18,  same,  except  5 
Mar.  1971;  18,  La  Pacifica,  4 km  NW  Canas,  14  Mar.  1972 
(P.A.  Opler;  UCB),  on  Inga  vera\  18,  Hacienda  Comelco,  24 
km  NW  Canas,  21  Mar.  1971  (E.R.  Heithaus;  LACM),  on 
Stachytarpheta  jamaicense,  0815;  18,  same  locality,  13  Mar. 
1971  (E.R.  Heithaus;  LACM),  on  Centrosoma  pubescens, 
0750. 

Epicharis  ( Epicharana ) rustic  a (Olivier) 

Figures  63,  67 

Apis  rustica  Olivier,  1789:64. 

Apis  hirtipes  Fabricius,  1793:325.  9. 

Epicharis  rustica:  F.  Smith,  1854:368. 

Centris  ( Epicharis ) rustica:  Friese,  1900b:253.  9 8. 

Centris  (Epicharis)  rustica  var.  /lava  Friese,  1900b:254.  8. 
NEW  SYNONYMY. 

Epicharis  rustica:  Moure,  1945a:295-296.  8 9 (syn.). 
Epicharis  (Epicharana)  rustica:  Michener,  1954:144  (distr.). 
Epicharis  ( Epicharis ) rustica:  Moure,  1 960b:  1 19-120  (syn.). 
Epicharis  (Epicharis)  flava:  Moure,  1960b:  120  (status). 

Moure  (1960b)  reexamined  the  type  female  of  Apis  hirtipes 
and  reaffirmed  its  traditional  place  in  the  synonymy  of  E. 
rustica.  He  concluded  “.  . . that  with  this  discovery,  the  true 
meaning  of  Epicharis  is  to  be  restored  as  in  my  revision  of 
Epicharis  (1945)  and  Epicharana  Michener,  1954,  with  the 
same  type  species  is  to  be  considered  a synonym.”  I have 
already  alluded  to  be  unavailability  of  Apis  rustica  to  be  the 
type-species  of  Epicharis. 


In  point  of  fact,  Moure’s  determination  is  inconclusive. 
Moure  synonymized  A.  hirtipes  under  “.  . . Epicharis  rustica 
as  interpreted  by  older  authors  and  Friese  in  his  Monographic 
der  Bienengattung  Centris  (s.  lat.).”  This  is  by  no  means  the 
same  as  placing  A.  hirtipes  in  synonymy  with  Olivier’s  species. 
Moure  admitted  that  he  had  been  unable  to  find  Olivier’s 
type.  In  truth  the  identity  of  E.  rustica  is  unknown  and  the 
current  interpretation  of  this  species  rests  upon  the  very  in- 
secure foundation  of  assumptions  made  over  a century  ago 
by  workers  whose  concepts  of  species  were  different  from 
those  of  the  present  and  who  may  not  actually  have  seen  the 
relevant  specimens. 

For  the  present,  it  would  seem  best  to  continue  to  accept 
the  traditional  interpretation  of  E.  rustica  and  its  appended 
synonymy,  as  published  by  Moure  (1960b).  The  alternative 
would  be  to  regard  Olivier’s  species  as  unidentifiable  and  to 
use  the  next  available  name  (A.  hirtipes ),  suffering  the  re- 
sultant nomenclatural  confusion.  I adhere  to  the  traditional 
understanding,  which  includes  the  unavailability  of  E.  rustica 
as  the  type  species  for  this  genus. 

This  is  a common  bee,  ranging  from  Costa  Rica  and  Pan- 
ama to  Brazil  and  Peru.  Friese’s  var.  flava,  with  whitish 
cinereous  pubescence  on  the  thorax,  was  thought  by  Moure 
( 1 960b)  to  be  probably  a good  species.  In  general,  specimens 
from  northeastern  South  America  have  almost  entirely  black 
thoracic  pubescence  and  would  correspond  to  the  “typical” 
form  of  E.  rustica.  Similar  dark  specimens  are  also  found  in 
Costa  Rica,  but  most  populations  have  considerable  pale 
pubescence  on  the  dorsum  and  sides  of  the  thorax.  There 
are,  however,  many  variants,  the  most  common  being  a form 
with  dark  mesepisternal  hairs,  those  of  the  mesoscutum  and 
scutellum  pale,  or  largely  so.  These  specimens  of  an  inter- 
mediate character  occupy  geographical  areas  between  the  two 
extremes,  as  would  be  expected  if  E.  rustica  and  E.  flava 
were  conspecific,  which  I believe  to  be  the  case. 

NEW  RECORDS 

COSTA  RICA,  GUANACASTE:  19  [Hacienda]  Comelco,  5 
Mar.  1971  (P.A.  Opler;  UCB),  on  Tabebuia  rosea.  HERE- 
DIA: 19,  Finca  La  Selva,  500  m elev.,  23  June  1979  (D.R. 
Perry;  LACM).  PUNTARENAS:  19,  Monteverde,  1400  m 
elev.,  17  Sept.  1982  (C.D.  Nagano  and  M.  Hayes;  LACM). 
PANAMA,  CANAL  ZONE:  19,  Barro  Colorado  Island,  15 
July  1980  (H.  Wolda;  ROUB);  19,  same,  except  1 Aug.  1980. 
CHIRIQUI:  399;  18,  Dolega,  15  Mar.  1980  (D.W.  Inouye; 
ROUB).  PANAMA:  399,  Arraijan.  16  Oct.  1980  (D.  Roubik; 
ROUB);  499,  Chihbre  Cave,  26  July  1966  (R.D.  Sage;  UCB); 
18,  San  Miguelito,  16  May  1974  (M.  Gonzalez;  LACM). 

Subgenus  Par  epicharis  Moure 

Parepicharis  Moure,  1945a:307-308.  Type-species:  “ Pare - 
picharis  zonata  (Smith,  1854 )”  = Epicharis  zonata  F. 
Smith,  1854;  monobasic  and  original  designation. 

DESCRIPTION 

Maxillary  palp  two-segmented,  second  segment  much  longer 
than  first,  more  than  five  times  longer  than  wide,  gradually 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  59 


narrower  distad;  lateral  ridges  of  clypeal  disc  weak;  malar 
space  linear,  eye  margin  nearly  contiguous  with  mandible 
base;  frontal  carina  ending  before  attaining  anterior  ocellus; 
occipital  margin  abruptly  rounded;  flagelliform  occipital  se- 
tae reaching  about  to  level  of  anterior  margin  of  tegula;  pos- 
terior margin  of  dorsal  face  of  scutellum  impressed;  meta- 
notum  vertical;  jugal  lobe  of  posterior  wing  about  as  long  as 
cubital  cell  and  one-half  as  long  as  vannal  lobe. 

Female.  Labrum  with  median  ridge  weak  or  absent;  outer 
face  of  mesobasitarsus  with  mixed  long,  coarse  plumose  setae 
and  long,  fine  plumose  hairs  on  anterior  half,  posterior  half 
with  long,  coarse  plumose  setae  only;  basitibial  plate  without 
secondary  plate;  metatibia  no  longer  than  metabasitarsus; 
third  and  fourth  terga  without  specialized  basal  areas;  mar- 
gins of  pygidial  plate,  in  dorsal  view,  nearly  straight,  apex 
broadly  truncate. 

Male.  Labrum  without  median  ridge;  first  flagellar  segment 
longer  than  scape  and  longer  than  following  two  segments 
combined;  ocellocular  distance  less  than  ocellar  diameter; 
procoxa  without  apicoventral  process;  mesosternal  protu- 
berances absent;  metatrochanter  and  metafemur  without 
ventral  seta  patch;  metatibia  without  carinate  posteroventral 
ridge;  metabasitarsus  with  or  without  anterior  carina-like 
ridge,  when  present  terminating  in  tooth  a little  beyond  mid- 
length; pygidial  plate  narrower  than  seventh  tergite,  margins 
cariniform,  apex  narrow,  rounded  or  truncate. 

Parepicharis  was  proposed  as  a monotypic  genus  for  Epi- 
charis  zonata.  Subsequently,  Moure  and  Seabra  (1959)  added 
E.  metatarsalis  to  Parepicharis.  Of  the  two  species,  only  E. 
metatarsalis  is  found  in  Central  America.  I have  seen  ma- 
terial of  E.  zonata  from  Guyana,  Brazil,  and  Bolivia  and 
there  are  records  of  the  species  from  Peru  and  Trinidad. 

KEY  TO  SPECIES  OF  PAREPICHARIS 

la.  Male,  antenna  13-segmented  and  ocellocular  distance 

less  than  diameter  of  anterior  ocellus  2 

b.  Female,  antenna  1 2-segmented  and  ocellocular  distance 

greater  than  diameter  of  anterior  ocellus  3 

2a.  Metabasitarsus  about  twice  as  long  as  broad  and  with 
distinct  tooth  on  anterior  margin  . . metatarsalis  Friese 
b.  Metabasitarsus  about  six  times  longer  than  broad  and 
without  tooth  on  anterior  margin  ....  zonata  F.  Smith 
3a.  First  abdominal  segment  blackish,  remaining  segments 
dull  ferruginous,  immaculate;  prepygidial  fimbria  con- 
sisting of  long,  closely  ranked,  plumose  hairs  

metatarsalis  Friese 

b.  Abdomen  brown,  with  conspicuous  yellow  maculations 
on  at  least  first  three  terga;  prepygidial  fimbria  weak, 
consisting  of  short,  discretely  separated,  short-plumose 

hairs  which  do  not  conceal  underlying  surface  

zonata  F.  Smith 

Epicharis  ( Parepicharis ) metatarsalis  Friese 

Epicharis  metatarsalis  Friese,  1899:40.  <5. 

Epicharis  phenacura  Cockerell,  1917:200.  6.  NEW  SYN- 
ONYMY. 


Epicharis  conura  Cockerell,  1917:200.  9.  NEW  SYNONY- 
MY. 

Epicharis  ( Parepicharis ) metatarsalis:  Moure  and  Seabra, 
1959:126  (distr.,  tax.). 

The  male  of  E.  metatarsalis  differs  from  that  of  E.  zonata 
most  obviously  by  the  metabasitarsus,  which  is  only  twice 
as  long  as  broad  and  with  a distinct  tooth  on  the  anterior 
margin  beyond  the  middle.  Males  of  E.  zonata  lack  a tooth 
on  the  metabasitarsus  and  the  segment  is  about  six  times 
longer  than  broad.  Females  of  E.  metatarsalis  have  the  first 
tergite  blackish  and  the  remaining  segments  dull  yellowish 
red;  in  E.  zonata  there  are  yellow  maculae,  of  variable  extent, 
on  the  first  three  tergites  which  usually  are  dark  brown  to 
blackish.  Both  sexes  of  E.  metatarsalis  are  larger  (25-27  mm 
versus  20-23  mm). 

Friese  (1900b)  recorded  males  of  this  species  from  St.  Pa- 
rime,  Venezuela  (type  locality),  and  “Darien  (=Colombia),” 
now  in  Panama.  Another  male  was  reported  by  Moure  and 
Seabra  (1959)  from  San  Jose,  Costa  Rica.  San  Carlos,  Costa 
Rica  is  the  type  locality  for  both  E.  phenacura  and  E.  conura. 
The  material  now  available  indicates  that  E.  phenacura  and 
E.  conura  are  the  opposite  sexes  of  one  species  and  that  the 
males  are  inseparable  from  E.  metatarsalis. 

NEW  RECORDS 

COSTA  RICA,  HEREDIA:  2399,  633,  Finca  La  Selva,  near 
Puerto  Viejo,  5 June-24  July  (D.R.  Perry;  LACM),  on  Hy- 
menolobium  sp.  (899,  288),  Dipteryx  panamensis  (1 199,  233), 
Tabebuia  sp.  (1<S),  Dussia  sp.  (13),  and  Byrsonima  sp.  (499). 

Subgenus  Hoplepicharis  Moure 

Hoplepicharis  Moure,  1945a:300-301 . Type-species:  ‘'Hop- 
lepicharis fasciata  (Lepeletier  & Serville,  1 828)”  = Epi- 
charis fasciata  Lepeletier  and  Serville,  1828;  original  des- 
ignation. 

Epicharis  subg.  Hoplepicharis:  Michener,  1954:145. 

DESCRIPTION 

Maxillary  palp  two-segmented,  second  segment  shorter  than 
first,  flattened;  lateral  ridges  of  clypeal  disc  strong;  malar 
space  distinct,  slightly  longer  than  minimum  thickness  of 
first  flagellar  segment;  frontal  carina  sharp  but  ending  well 
in  front  of  anterior  ocellus;  occipital  margin  abruptly  round- 
ed; flagelliform  occipital  setae  extending  nearly  to  level  of 
posterior  tegular  margin;  posterior  margin  of  dorsal  face  of 
scutellum  not  impressed;  metanotum  bifaced,  juncture  sharply 
angulate  to  crested;  jugal  lobe  of  posterior  wing  about  as  long 
as  cubital  cell  and  about  one-half  as  long  as  vannal  lobe. 

Female.  Labrum  with  weak  median  ridge;  outer  face  of 
mesobasitarsus,  on  anterior  half,  with  long,  coarse,  simple 
setae  and  shorter,  fine,  plumose  hairs,  posterior  half  with 
sparse  long,  coarse,  simple  setae  and  scattered  short,  fine, 
plumose  hairs;  secondary  basitibial  plate  present;  metatibia 
a little  longer  than  metabasitarsus;  third  and  fourth  terga 
without  specialized  basal  areas;  margins  of  pygidial  plate 


60  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


nearly  straight  and  strongly  convergent  to  narrowly  truncate 
apex. 

Male.  Labrum  without  median  ridge;  first  flagellar  segment 
shorter  than  either  scape  or  second  flagellar  segment;  ocel- 
locular  distance  greater  than  diameter  of  anterior  ocellus; 
procoxa  with  short,  inconspicuous  apicoventral  process; 
mesosternum  without  tubercles;  metatrochanter  and  meta- 
femur without  ventral  seta  patch;  metatibia  without  pos- 
teroventral  ridge;  metabasitarsus  with  anterior  carinate  ridge 
terminating  in  sharp  tooth  basad  of  midlength;  pygidial  plate 
broad  and  indistinct,  weakly  bilobate  at  apex. 

This  is  a small  group,  with  two  species  known  from  South 
America  and  two  in  Central  America. 

KEY  TO  CENTRAL  AMERICAN  HOPLEPICHARIS 

a.  Second  abdominal  tergite  with  basal  yellow  band,  nar- 

rowed across  middle,  followed  by  narrow  blackish  brown 
band;  most  of  remainder  of  this  segment,  and  all  of  fol- 
lowing segments,  ferruginous  (except  short,  yellow  lon- 
gitudinal stripe  at  side  of  third  segment)  

lunulata  Mocsary 

b.  Similar,  but  second  tergite,  beyond  yellow  basal  band, 

and  all  of  following  segments,  blackish  (except  short,  yel- 
low longitudinal  stripe  at  side  of  third  segment) 

monozona  Mocsary 

Epicharis  ( Hoplepicharis ) lunulata  Mocsary 

Figure  64 

Epicharis  lunulata  Mocsary,  1898:499.  <5  9. 

Hoplepicharis  lunulata:  Moure,  1945a:302  (distr.,  syn.). 
Epicharis  ( Hoplepicharis ) lunulata:  Michener,  1954:145 
(distr.). 

This  attractive  Central  American  species  appears  to  be  less 
common  than  the  superficially  similar  E.  e/egans.  Its  range 
extends  from  Mexico  to  Panama. 

NEW  RECORDS 

MEXICO,  CHIAPAS:  19,  Simojovel,  18-31  July  1958  (J.A. 
Chemsak;  UCB).  JALISCO:  19,  Estacion  Biologia  UNAM, 
Chamela,  1 1 Sept.  1981  (S.H.  Bullock;  LACM),  on  Psidium 
sartorianum.  NAYARIT:  299,  16  mi.  NW  Tepic,  19  July  1953 
(Univ.  Kans.  Mex.  Exped.;  UKAN).  OAXACA:  19,  6 mi.  W 
Zanatepec,  150  ft.  elev.,  9 July  1953  (Univ.  Kans.  Mex. 
Exped.;  UKAN),  on  Malpighia  mexicana\  299,  5 mi.  NW 
Totolapan,  3800  ft.  elev.,  6 July  1953  (Univ.  Kans.  Mex. 
Exped.;  UKAN),  on  Malpighia  mexicana.  VERA  CRUZ:  19, 
13,  Cordoba,  1-10  Sept.  (9),  1 Oct.  1964  (<3)  (LACM).  GUA- 
TEMALA: 19,  Cayuga,  Nov.  (Schaus  and  Barnes;  USNM); 
19,  “F.4694”  (UKAN).  HONDURAS:  13,  Tegucigalpa,  12 
May  1981  (F.J.  Dyer;  USNM).  COSTA  RICA,  GUANA- 
CASTE:  499,  13,  Hacienda  Comelco,  8 km  NW  Bagaces  and 
24  km  NW  Canas,  dates  between  7 Mar.  and  14  Oct.  (E.R. 
Heithaus  [19],  P.A.  Opler  [399,  13];  LACM,  UCB),  on  Byr- 
sonima  sp.  (19),  Stachytarphe  frantzii  (19),  Petastoma  patel- 
liferum  (3)  and  Bignonaceae,  0700  (19);  19,  near  Turin 
(10°20'N,  84°50'W),  1 Feb.  1960  (C.W.  Palmer;  UKAN). 


PUNTARENAS:  19,  1 mi.  ESE  jet.  Rio  Canas  and  Hwy  2, 
1000  ft.  elev.,  23  July  1965  (R.D.  Sage  and  S.J.  Arnold;  UCB), 
on  Bixa  orel/ana,  1000-1 100.  SAN  JOSE:  13,  4 km  E San 
Ignacio  de  Acosta,  4000  ft.  elev.,  8 July  1 963  (C.D.  Michener 
et  ah;  UKAN);  19,  Playon,  8 km  N Parrita,  30  ft.  elev.,  14- 
19  Aug.  1962  (C.D.  Michener  and  A.  Wille;  UKAN);  13, 
San  Jose,  25  July  1913  (UKAN). 

Epicharis  ( Hoplepicharis ) monozona  Mocsary 

Epicharis  monozona  Mocsary,  1898:498.  9. 

Hoplepicharis  monozona:  Moure,  1945a:392  (distr.,  tax.). 
Epicharis  ( Hoplepicharis ) monozona:  Michener,  1954:145 
(distr.). 

This  species  was  based  on  a female  from  an  unspecified  Pan- 
amanian locality.  Moure  (1945a)  recorded  another  female 
from  Muzo,  Rio  Cantinero,  Colombia,  and  Michener  ( 1954) 
reported  two  additional  Panamanian  specimens,  both  from 
the  Canal  Zone:  Las  Cruces  trail,  near  Corozal,  and  Fort 
Clayton. 

The  male  of  E.  monozona  has  not  been  previously  re- 
ported. It  is  separable  from  that  of  E.  lunulata  by  the  black, 
rather  than  ferruginous,  abdomen.  Aside  from  the  differences 
in  color,  E.  monozona  and  E.  lunulata  appear  to  be  identical 
and  I suspect  they  will  ultimately  be  found  to  be  conspecific. 

NEW  RECORDS 

PANAMA,  PANAMA:  399,  Arraijan,  16  Oct.  1980  (D.  Rou- 
bik;  LACM,  ROUB);  19,  Chepo,  15  km  E Carti,  8 June  (D. 
Roubik;  ROUB);  13,  Cerro  Cantpana,  13  Apr.  1960  (W.J. 
Hanson;  UKAN);  19,  Cerro  Azul,  N of  Tocumen,  15  June 
1958  (W.J.  Hanson;  UKAN).  CANAL  ZONE:  19,  Pipeline 
Road,  Gamboa,  12  Jan.  1980  (D.  Roubik;  ROUB);  19,  same, 
except  10  June  1980;  19,  7 33,  Pipeline  Road,  20  Sept.,  6 Oct. 
1979  (K.  Steiner;  UCD),  on  Drymonia  serrulata:  599,  13, 
Gamboa,  28  Sept.  1979,  same  collector  and  host;  13,  Barro 
Colorado  Island,  15  July  1958  (W.J.  Hanson;  UKAN);  333, 
same  locality,  15  Sept.,  12  Oct.,  28  Oct.  1979  (K.  Steiner; 
UCD),  on  D.  serrulata.  COLON:  19,  Portobello,  30  Sept. 
1979  (K.  Steiner;  UCD),  on  D.  serrulata. 

Subgenus  Epicharoides  Radoszkowski 

Epicharoides  Radoszkowski,  1884:20.  Type-species:  (Epi- 
charoides bipunctatus  Radoszkowski,  1884 ) = Epicharis 
maculata  F.  Smith,  1874;  monobasic  and  original  desig- 
nation. 

Epicharoides:  Moure,  1945a:309-310. 

Epicharis  subg.  Epicharoides:  Michener,  1954:144. 

DESCRIPTION 

Maxillary  palp  three-segmented,  second  segment  longer  than 
first,  third  segment  narrower  and  shorter  than  second;  lateral 
ridges  of  clypeal  disc  strong;  malar  space  distinct,  about  as 
long  as  minimum  thickness  of  first  flagellar  segment;  frontal 
carina  sharp,  nearly  reaching  anterior  ocellus;  occipital  mar- 
gin slightly  compressed  but  not  ridge-like;  flagelliform  oc- 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  61 


Figures  77-80.  Epicharis  (Epicharoides)  albofasciata,  male  seventh  and  eighth  stemites  and  genitalia  (ventral  and  dorsal  views).  Scale  line  = 
1.00  mm. 


cipital  setae  short  and  curved  laterad,  not  extending  beyond 
anterior  margin  of  mesoscutum;  posterior  margin  of  dorsal 
face  of  scutellum  not  impressed;  metanotum  wholly  vertical; 
jugal  lobe  of  posterior  wing  about  as  long  as  cubital  cell  and 
about  one-half  as  long  as  vannal  lobe. 


Female.  Labrum  without  median  ridge;  mesobasitarsis  ex- 
ternally with  evenly  distributed  long  coarse  setae  (some  plu- 
mose) and  short,  fine,  long-plumose  hairs;  secondary  basi- 
tibial  plate  absent;  metatibia  no  longer  than  metabasitarsus; 
third  and  fourth  terga  without  specialized  basal  areas;  mar- 


62  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


gins  of  pygidial  plate,  in  dorsal  view,  slightly  concave,  apex 
narrowly  truncate,  disc  depressed. 

Male.  Labrum  without  median  ridge;  scape  unusually  ro- 
bust, slightly  longer  than  wide,  first  flagellar  segment  shorter 
than  either  scape  or  second  flagellar  segment;  ocellocular 
distance  greater  than  diameter  of  anterior  ocellus;  procoxa 
without  apicoventral  process;  mesosternal  tubercles  absent; 
metatrochanter  and  metafemur  without  ventral  seta  patch; 
metatibia  without  posteroventral  ridge;  metabasitarsus  with- 
out carinate  anterior  ridge;  pygidial  plate  about  one-half  as 
broad  as  seventh  tergite,  sharply  margined,  apex  narrowly 
rounded. 

Moure  (1945a)  recognized  three  species  in  this  subgenus, 
and  a fourth  was  added  by  Moure  and  Seabra  (1959).  Two 
of  these  species  are  present  in  Central  America  and  two  are 
confined  to  South  America.  The  two  Central  American  species 
are  black,  with  abundant,  very  variable  yellow  markings. 

KEY  TO  CENTRAL  AMERICAN  EPICHAROIDES 

a.  Male  clypeus  with  irregular  black  blotch  on  disc;  female 

with  basal  yellow  band  of  second  tergite  broadly  inter- 
rupted, but  if  complete,  it  is  much  shorter  in  middle  than 
black  band  following  it  and  basal  bands  of  following  terga 
are  broadly  interrupted albofasciata  F.  Smith 

b.  Male  clypeus  without  black  blotch  on  disc;  female  with 
basal  yellow  band  of  second  tergum  as  long  in  middle  as 
black  band  following  it  or,  if  a little  shorter,  third  and 
fourth  terga  with  complete  transverse  yellow  fasciae  . . . 

maculata  F.  Smith 

Epicharis  {Epicharoides)  albofasciata  F.  Smith 

Figures  77-80 

Epicharis  albofasciata  F.  Smith,  1874:321.  <3. 

Epicharis  maculata  var.  nigroclypeata  Friese,  1899:40. 
Epicharoides  albofasciata:  Moure,  1945a:310  (syn.,  distr.). 
Epicharis  (Epicharoides)  maculata:  Michener,  1954:145  (in 
part). 

This  species  and  E.  maculata  are  so  similar  as  to  be  virtually 
inseparable  and  records  from  the  literature  under  either  name 
must  be  treated  with  caution.  There  are,  however,  consistent 
differences  in  the  male  terminalia  (compare  Figs.  77-80  with 
Figs.  81-84). 

Females  of  the  two  species  are  especially  difficult  to  sep- 
arate. Those  of  E.  albofasciata  have  either  lateral  spots  on 
the  second  tergite  or  a subbasal  fascia  which  is  shorter  than 
the  black  area  following  the  band;  the  third  and  fourth  tergites 
apparently  never  possess  entire  transverse  subbasal  bands, 
though  the  margins  of  the  segments  may  be  broadly  ferru- 
ginous. On  the  other  hand,  females  of  E.  maculata  possess 
a subbasal  band  on  the  second  segment  which  is  usually, 
though  not  always,  at  least  as  long  as  the  dark  band  following 
it;  a shorter  transverse  band,  often  attenuated  in  the  middle, 
is  present  on  the  third  tergite;  a transverse  band  is  also  present 
on  the  fourth  segment,  usually  longer  than  that  of  the  third. 

Aside  from  the  differences  in  genitalic  structures,  males  of 
E.  albofasciata  are  recognizable  by  the  presence  of  a black 


blotch  on  the  the  clypeal  disc,  the  very  short  fasciae  of  the 
second  to  fourth  tergites  (evanescent  or  absent  on  third  and 
often  on  fourth  as  well)  and  the  pronotum  black,  except  two 
widely  separated  spots  on  the  collar. 

From  Central  America,  I have  seen  material  of  E.  albo- 
fasciata only  from  Costa  Rica  and  Panama.  At  least  some  of 
the  specimens  from  Panama  recorded  by  Michener  (1954) 
as  E.  maculata  are  this  species;  his  figures  141-143  are  based 
on  E.  albofasciata. 

NEW  RECORDS 

COSTA  RICA,  GUANACASTE:  299,  Hacienda  Comelco,  24 
km  NW  Canas,  6-13  Mar.  1972  (E.R.  Hcithaus;  LACM),  1 
on  Securidaca  tenuifolia ; 19,  Hacienda  Comelco,  8 km  NW 
Bagaces,  5 Mar.  1971  (P.A.  Opler;  UCB),  on  Delbergia  re- 
teusa\  1<3,  same  locality,  28-31  Jan.  1972  (P.A.  Opler;  UCB), 
on  Securidaca  sylvestris\  13,  Finca  La  Pacifica,  4 mi.  NW 
Canas,  10  July  1971  (P.A.  Opler;  UCB),  on  Petastoma  pa- 
telliferum ; 19,  Liberia,  15-16  Feb.  1972  (P.A.  Opler;  UCB), 
on  Andira  inermis.  HEREDIA:  599,  1633,  Finca  La  Selva, 
near  Puerto  Viejo,  6 May  to  18  June  (D.R.  Perry;  LACM), 
on  Dipteryx  panamensis  (399,  833),  Vochysia  sp.  (13),  Byr- 
sonima  sp.  (19),  Dussia  sp.  (633),  and  Hymenolobium  sp. 
(13).  PANAMA,  PANAMA:  19,  24  Mar.  1980  (D.  Roubik; 
ROUB);  499,  same,  except  2 Apr.  1980;  299,  same  except  13 
Apr.  1980;  19,  Curundu,  3 1 Mar.  1981  (D.  Roubik;  ROUB). 
CANAL  ZONE:  19,  Barro  Colorado  Island,  2 June  1981  (H. 
Wolda;  ROUB). 

Epicharis  ( Epicharoides ) maculata  F.  Smith 

Figures  8 1-84 

Epicharis  maculata  F.  Smith,  1874:320.  9. 

Epicharoides  bipunctatus  Radoszkowski,  1884:20. 

Centris  (Epicharis)  variabilis  Friese,  1 900b: 3 5 1 . (New  name 
for  C.  maculata  F.  Smith,  not  C.  maculata  (Lepeletier.) 
Epicharoides  maculata:  Moure,  1945a:310  (syn.,  distr.). 
Epicharis  (Epicharoides)  maculata:  Michener,  1954: 145  (syn., 
distr.)  (in  part). 

This  species  was  described  from  Oaxaca,  Mexico,  and  ranges 
south  to  Venezuela  and  Colombia.  In  Central  America  it  is 
much  more  common  than  E.  albofasciata.  a primarily  South 
American  species. 

NEW  RECORDS 

GUATEMALA:  19,  Secanquim,  June  1984  (USNM);  19, 
Quirigua,  Aug.  (Schaus  and  Barnes;  USNM).  MEXICO,  OA- 
XACA: 299,  Salina  Cruz,  10  Aug.  1964  (E.  Fisher  and  D. 
Verity;  LACM).  QUINTANA  ROO:  19,  3 33,  8-14  May  1963 
(E.C.  Welling;  LACM).  SINALOA:  19,  5 mi.  N Mazatlan,  29 
July  1 973  (J.A.  Chemsak,  E.G.  Linsley,  A.E.  and  M.M.  Mich- 
elbacher;  UCB),  on  Turnera  diffusa.  VERA  CRUZ:  599,  Te- 
colulla,  19  June  1951  (P.D.  Hurd;  UCB).  YUCATAN:  19, 
Piste,  July  1967  (E.C.  Welling;  LACM).  COSTA  RICA, 
GUANACASTE:  19,  Liberia,  15-16  Feb.  1972  (P.A.  Opler; 
UCB),  on  Andira  inermis\  299,  Finca  La  Pacifica,  4 km  NW 
Canas,  17-20  Nov.  1972  (P.A.  Opler;  UCB),  on  “pink  malp. 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  63 


Figures  81-84.  Epicharis  (Epicharoides)  maculata,  male  seventh  and  eighth  sternites  and  genitalia  (ventral  and  dorsal  views).  Scale  line  = 
1 .00  mm. 


vine”;  19,  same  except  30  Nov.  1 972,  on  Bixa  orellana\  1 133, 
Hacienda  Comelco,  24  km  NW  Canas,  13-14  Mar.  1971 
(E.R.  Heithaus;  LACM),  on  Secundaca  tenuifo!ia\  333,  Ha- 
cienda Comelco,  8 km  NW  Bagaces,  28-3  1 Jan.,  6 Mar.  1971 


(P.  A.  Opler;  UCB),  on  Secridaca  sylvestns.  HEREDIA:  1 099, 
7133,  Ftnca  La  Selva,  near  Puerto  Viejo,  25  Apr.  to  19  June 
(D.R.  Perry;  LACM),  on  Vochysia  sp.  (533),  Byrsonima  sp. 
(599),  Diptervx panamensis  (599,  6033),  Dussia  sp.  (333),  and 


64  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


Hymenolobium  sp.  (1<5).  PANAMA,  PANAMA:  2<3<5,  Capira, 
Cerro  Campana,  12  Aug.  1980  (D.  Roubik;  ROUB);  1<3,  15 
km  E Chepo,  Llano  Carti  Rd.,  22  Feb.  1980  (D.  Roubik; 
ROUB). 

Subgenus  Epicharitides  Moure 

Epicharitides  Moure,  1945a:  311.  Type-species:  “Epichari- 
tides cockerelli  (Friese,  1900)”  = Epicharis  cockerelli  Fnese, 

1900a;  original  designation. 

DESCRIPTION 

Maxillary  palp  three-segmented,  second  segment  broad,  longer 
than  first,  third  segment  narrower  and  a little  shorter  than 
second;  lateral  ridges  of  clypeal  disc  strong;  malar  space  dis- 
tinct, but  shorter  than  minimum  thickness  of  first  flagellar 
segment;  frontal  carina  sharp  but  short,  ending  in  front  of 
anterior  ocellus  by  much  more  than  diameter  of  anterior 
ocellus;  occipital  margin  compressed  and  ridge-like  or  crest- 
ed; flagelliform  occipital  setae  short  and  curved  laterad  before 
reaching  anterior  margin  of  mesoscutum;  posterior  margin 
of  dorsal  face  of  scutellum  not  impressed;  metanotum  wholly 
vertical,  jugal  lobe  of  posterior  wing  shorter  than  cubital  cell 
and  less  than  one-half  as  long  as  vannal  lobe. 

Female.  Labrum  without  median  ridge;  outer  face  of  me- 
sobasitarus  with  mixed  long,  coarse,  simple  setae  and  short, 
fine,  long-plumose  hairs  on  anterior  half,  posterior  half  with 
sparse  long,  coarse,  simple  setae  only;  secondary  basitibial 
plate  absen;  metatibia  no  longer  than  metabasitarsus;  grad- 
ulus  of  third  and  fourth  terga,  in  middle,  broadly  deflected 
apicad  and  area  on  either  side  depressed  and  covered  with 
dense  mat  of  very  short,  plumose  hairs;  margins  of  pygidial 
plate,  in  dorsal  view,  nearly  straight,  apex  narrowly  truncate, 
disc  depressed. 

Male.  Labrum  without  median  ridge;  first  flagellar  segment 
shorter  than  either  scape  or  second  flagellar  segment;  ocel- 
locular  distance  greater  than  diameter  of  anterior  ocellus; 
procoxa  without  apicoventral  process;  mesosternal  tubercles 
absent;  metatrochanter  and  metafemur  without  ventral  seta 
patch;  metatibia  without  posteroventral  ridge;  metabasitar- 
sus without  carinate  ridge  on  anterior  margin;  pygidial  plate 
less  than  one-half  as  wide  as  seventh  tergite,  margins  sharp, 
apex  narrowly  truncate. 

Moure  (1945b)  recognized  five  species  in  this  subgenus 
and  gave  a key  for  their  separation.  Two  additional  species 
were  described  from  Brazil  by  Moure  and  Seabra  ( 1959).  All 
the  included  species  were  known,  at  that  time,  only  from 
South  America,  especially  Brazil.  One  species  is  now  known 
to  occur  in  Central  America. 

Epicharis  ( Epicharitides ) species 

Two  females  from  Cerro  Campana,  Panama  Province,  Pan- 
ama, 9 June  1960  (W.J.  Hanson;  UKAN)  possibly  represent 
an  undescribed  species.  In  the  key  to  species  of  Epicharitides 
by  Moure  (1945b)  they  fail  at  couplet  7,  failing  to  agree  with 
the  characteristics  cited  for  either  E.  obscura  Friese  or  E. 
duckei  Friese.  The  abdomen  is  black  ventrally  as  well  as 


dorsally,  there  is  a narrow  transverse  lateral  stripe  on  each 
side  of  the  dorsal  face  of  the  first  tergum  and  the  second 
tergum  has  a broad  basal  yellow  band,  slightly  expanded  on 
either  side.  The  distal  margin  of  this  band  is  broadly  curved 
inward,  but  in  the  middle  of  the  segment,  the  band  is  one- 
half  as  long  as  the  segment.  The  face  is  wholly  black  except 
for  the  yellow  basal  portion  of  the  labrum  and  a minute 
median  spot  on  the  lower  paraocular  area. 

These  specimens  may  represent  a previously  undescribed 
species,  but  in  the  absence  of  associated  males,  no  nomen- 
clatural  action  is  possible  at  this  time. 

These  specimens  differ  from  E.  duckei  in  the  color  pattern 
of  the  abdominal  terga.  The  first  tergite  bears  a short,  sub- 
lateral  yellow  stripe  and  the  second  is  yellow  across  the  entire 
base,  with  the  distal  one-third  blackish,  the  yellow  band  a 
little  shorter  in  the  middle  one-half.  The  remainder  of  the 
abdomen  is  blackish.  Pubescence  is  wholly  dark,  except  the 
pale  scopa  and  some  pale  hairs  around  the  pronotal  lobe. 
Females  of  E.  obscura  agree  with  the  Panamanian  specimens 
in  having  the  pubescence  largely  dark,  but  the  first  tergum 
is  immaculate  and  the  second  to  fourth  or  fifth  terga  are 
maculate  only  at  the  sides. 

Superficially,  these  females  resemble  unusually  dark  in- 
dividuals of  E.  maculata,  but  may  be  readily  separated  by 
the  shorter  jugal  lobe  of  the  hind  wing,  by  the  characteristic 
shape  of  the  metabasitarsus  and  by  the  presence  of  the  mod- 
ified areas  on  the  abdominal  terga.  These  differences  will  also 
distinguish  this  bee  from  E.  albofasciata. 

SYNONYMIC  LIST  OF  NORTH  AND  CENTRAL 
AMERICAN  CENTRIDINI 

Centris  Fabricius 

Subgenus  Xerocentris  Snelling,  1974.  Type-species:  C.  cali- 
fornica  Timberlake,  1940. 

01.  californica  Timberlake,  1940.  Calif.,  Nev. 

02.  griseola  Snelling,  n.  sp.  Mexico  (Guerrero). 

03.  hoffmanseggiae  Cockerell,  1897.  N.  Mex.,  Ariz.,  Calif. 

= davidsoni  Cockerell,  1904. 

04.  pallida  W.  Fox,  1899.  Southwestern  U.S.,  northwestern 

Mexico. 

= callognatha  Cockerell,  1923. 

= trichosoma  Cockerell,  1923. 

05.  rhodomelas  Timberlake,  1940.  Calif. 

06.  tiburonensis Cockerell,  1923.  Calif.,  Ariz.,  Son.,  B.  Calif. 

07.  vanduzeei  Cockerell,  1923.  Mexico  (B.  Calif.,  B.  Calif. 

Sur). 

Subgenus  Paracentris  Cameron,  1902.  Type-species:  C.  ful- 
vohirta  (Cameron,  1902). 

= Penthemisia  Moure,  1950.  Type-species:  C.  chilensis 
Spinola,  1851. 

= Trichocentris  Snelling,  1956.  Type-species:  C.  rho- 
doleuca  Cockerell,  1923. 

08.  angustifrons  Snelling,  1966.  Ariz.,  Son. 

09.  aterrima  F.  Smith,  1854.  Ariz.  to  central  Mexico. 

1 0.  atripes  Mocsary,  1 899.  Southwestern  U.S.  to  Costa  Rica. 
= atriventris  W.  Fox,  1899. 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  65 


= limbata  Friese,  1899.  N.  SYN. 

= Foxi  Friese,  1900b. 

11.  caesalpiniae  Cockerell,  1897.  Tex.  to  Ariz.,  northern 

Mexico. 

= morsei  Cockerell,  1897. 

= marginata  W.  Fox,  1899. 

12.  cockerelli  W.  Fox,  1899.  Southwestern  U.S.,  northern 

Mexico. 

= resoluta  Cockerell,  1923.  N.  SYN. 

13.  ectypha  Snelling,  1974.  B.  Calif. 

14.  ferrisi  Cockerell,  1924.  B.  Calif.  Sur. 

15.  fisheri  Snelling,  1974.  B.  Calif,  B.  Calif.  Sur. 

16.  harbisoni  Snelling,  1974.  B.  Calif 

17.  laevibullata  Snelling,  1966.  Central  Mexico. 

18.  lanosa  Cresson,  1872.  Fla.  to  Tex.,  Okla.,  Kans. 

= subhyalina  W.  Fox,  1899. 

= birkmanii  Friese,  1900a. 

19.  mexicana  F.  Smith,  1854.  Texas  to  Ariz.,  south  to  Oa- 

xaca, Mexico. 

20.  nigrocaerulea  F.  Smith,  1874.  Central  Mexico  to  Pan- 

ama. 

= clypeata  Friese,  1899.  Preoccupied.  N.  SYN. 

= anthracina  Snelling,  1966.  N.  SYN. 

21.  rhodopus  Cockerell,  1 897.  Southwestern  U.S.,  northern 

Mexico. 

= pulchrior  Cockerell,  1900. 

= rhodoleuca  Cockerell.  1923. 

22.  zacateca  Snelling,  1966.  Ariz.,  N.  Mex.,  central  Mexico. 

Subgenus  Xanthemisia  Moure,  1945b.  Type-species:  C.  bi- 
color Lepeletier,  1841. 

23.  caro/ae  Snelling,  1966.  Mexico  to  Costa  Rica. 

24.  lutea  Friese,  1899.  Central  Mexico  to  Panama;  South 

America. 

25.  rubella  F.  Smith,  1854.  Panama;  South  America. 

Subgenus  Exa/locentris  Snelling,  1974.  Type-species:  C.  an- 
omala  Snelling,  1966. 

26.  anomala  Snelling,  1966.  Central  Mexico. 

Subgenus  Acritocentris  Snelling,  1974.  Type-species:  C.  ru- 
thannae  Snelling,  1966. 

27.  agameta  Snelling,  1974.  Northeastern  and  central  Mex- 

ico. 

28.  a/biceps  Friese,  1899.  Northeastern  and  central  Mexico. 
= strawi  Snelling,  1966.  N.  SYN. 

29.  ruthannae  Snelling,  1966.  Ariz.,  Son. 

30.  satana  Snelling,  n.  sp.  Southern  Arizona  to  central  Mex- 

ico. 

Subgenus  Centris  Fabncius,  1804.  Type-species:  C.  hae- 
morrhoidalis  (Fabricius,  1775). 

= Hemisia  Klug,  1807.  Type-species:  C.  haemorrhoi- 
dalis  (Fabricius,  1775). 

= Cyanocentris  Friese,  1900b.  Type-species:  C.  versi- 
color (Fabricius,  1775). 

= Poecilocentris  Friese,  1900b.  Type-species:  C.fascia- 
tella  Friese,  1 900b. 

31.  adanae  Cockerell.  1949.  Southern  Mexico  to  Panama. 

32.  aethiocesta,  Snelling,  n.  sp.  El  Salvador  to  Panama. 


33.  aethyctera  Snelling,  1974.  Southern  Mexico  to  Panama. 

34.  decolorata  Lepeletier,  1841.  Texas  to  Panama;  Carib- 

bean; northern  South  America. 

= obscuriventris  Friese,  1899? 

35.  eisenii  W.  Fox,  1899.  Arizona  to  Panama. 

36.  errans  W.  Fox,  1899.  Florida. 

37.  flavifrons  (Fabricius,  1775)  Central  America;  South 

America. 

= brasiliana  Christ,  1791. 

= citrotaemata  Gribodo,  1894.  N.  SYN. 

= nigritula  Friese,  1899.  N.  SYN. 

= rufescens  Friese,  1899.  N.  SYN. 

38.  jlavo/asciata  Friese,  1900a.  Central  America,  northern 

South  America. 

39.  inermis  Friese,  1899.  Central  America,  northern  South 

America. 

= segregata  Crawford,  1906.  N.  SYN. 

= gualanensis  Cockerell,  1912. 

= robusta  Cockerell,  1949.  N.  SYN. 

= pallidifrons  Cockerell,  1949. 

40.  meaculpa  Snelling,  n.  name.  Eastern  Mexico. 

= erubescens  Snelling,  1974.  Preoccupied. 

41.  obscurior  Michener,  1954.  Southern  Mexico  to  Panama. 

Subgenus  Ptilocentris  Snelling,  n.  subg.  Type-species:  C. /es- 
tiva F.  Smith,  1854. 

42.  /estiva  F.  Smith,  1854.  Costa  Rica  to  Venezuela  and 

Peru. 

= chlorura  Cockerell,  1919.  N.  SYN. 

Subgenus  Melanocentris  Friese,  1 900b.  Type-species:  C.  atra 
Friese,  1899. 

43.  agilis  F.  Smith,  1874.  Mexico  to  Honduras. 

= ignita  F.  Smith,  1874.  N.  SYN. 

= bakeri  Friese,  1912.  N.  SYN. 

= bakerella  Friese,  1913.  N.  SYN. 

= Epicharis  cisnerosi  Cockerell.  1949.  N.  SYN. 

44.  agiloides  Snelling,  n.  sp.  Mexico  to  Costa  Rica. 

45.  fl avilabris  Mocsary,  1899.  Costa  Rica  to  South  America. 
= boliviensis  Mocsary,  1899. 

46.  fusciventris  Mocsary,  1899.  Costa  Rica  to  South  Amer- 

ica. 

= scutellata  Friese,  1900b.  N.  SYN. 

47.  gelida  Snelling,  n.  sp.  Mexico,  Guatemala. 

48.  obso/eta  Lepeletier,  1841.  Mexico  to  South  America. 

= melanochlaena  F.  Smith,  1874.  N.  SYN. 

= Epicharis  zamoranensis  Cockerell,  1949.  N.  SYN. 

49.  plumipes  F.  Smith,  1854.  Costa  Rica  to  South  America. 

50.  sericea  Friese,  1899.  Central  Mexico. 

Subgenus  Trachina  Klug,  1810.  Type-species:  C.  longimana 
Fabricius,  1804. 

= Paremisia  Moure,  1945b.  Type-species:  C.  lineo/ata 
Lepeletier,  1841. 

51.  dentata  F.  Smith,  1854.  Southern  Mexico  to  South 

America. 

= proxima  Friese,  1900a. 

52.  eurypatana  Snelling,  n.  sp.  Mexico  (Jalisco). 

53 . fuse  at  a Lepeletier,  1841.  Southern  Mexico  to  South 

America. 


66  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


54.  heithausi  Snelling,  1974.  Guatemala  to  Costa  Rica. 

55.  labiata  Friese,  1904.  Mexico  to  Costa  Rica. 

= schwarzi  Cockerell,  1919.  N.  SYN. 

56.  longimana  Fabricius,  1804.  Nicaragua  to  South  Amer- 

ica. 

= personata  F.  Smith,  1874. 

57.  similis  (Fabricius,  1804).  Costa  Rica  to  northern  South 

America. 

= lineolata  Lepeletier,  1841. 

58.  vidua  Mocsary,  1899.  Belize  to  Panama. 

59.  xochipillii  Snelling,  n.  sp.  Mexico  (Oaxaca). 

Subgenus  Hemisiella  Moure,  1945b.  Type-species:  C.  lanipes 
(Fabricius,  1775). 

60.  dichrootricha  Moure,  1945b.  Panama;  South  America. 

6 1 . nitida  F.  Smith,  1 874.  Mexico  to  Panama;  northern  South 

America. 

= confinis  Perez,  1905.  N.  SYN. 

62.  transversa  Perez,  1905. 

= ruae  Cockerell,  1949.  N.  SYN. 

63.  trigonoides  Lepeletier,  1841.  Southern  Mexico  to  South 

America. 

= dentipes  F.  Smith,  1874.  N.  SYN. 

= hoplopoda  Moure,  1943. 

= rufomaculata  Cockerell,  1949.  N.  SYN. 

= subtarsata  Cockerell,  1949.  N.  SYN. 

64.  vittata  Lepeletier,  1841.  Mexico  to  South  America. 

= montezuma  Cresson,  1879. 

= breviceps  Friese,  1899. 

= friesei  Crawford,  1906.  Preoccupied. 

= costaricensis  Crawford,  1907. 

= erubescens  Friese,  1925.  N.  SYN. 

Subgenus  Heterocentris  Cockerell,  1899.  Type-species:  C. 
cornuta  Cresson,  1865. 

65.  analis  (Fabricius,  1804).  Mexico  to  South  America. 

= totonaca  Cresson,  1879. 

= otomita  Cresson,  1879. 

= minuta  Mocsary,  1899. 

= simplex  Friese,  1899. 

= durantae  Cockerell,  1949. 

= petreae  Cockerell,  1949. 

= petreae  var.  rufopicta  Cockerell,  1949. 

66.  bicornuta  Mocsary,  1899.  Mexico  to  South  America. 

67.  difformis  F.  Smith,  1854.  Costa  Rica;  South  America. 

68.  labrosa  Friese,  1899.  Costa  Rica  to  South  America. 

= triangulifera  Cockerell,  1949.  N.  SYN. 

Pti  lot  opus  Klug 

Ptilotopus  Klug,  1810.  Type-species:  P.  americanus  Klug, 
1810. 

69.  zonatus  Mocsary,  1899.  Panama. 

= pandora  Friese,  1900b. 

Epicharis  Klug 

Subgenus  Epicharana  Michener,  1954.  Type-species:  E.  rus- 
tica  (Olivier,  1789). 

70.  angulosa  Snelling,  n.  sp.  Costa  Rica. 


7 1 . bova  Snelling,  n.  sp.  Costa  Rica,  Panama. 

72.  elegans  F.  Smith,  1861.  Mexico  to  Costa  Rica. 

= salazari  Cockerell,  1949.  N.  SYN. 

73.  rustica  (Olivier,  1789).  Costa  Rica  to  South  America. 

= flava  Friese,  1900b.  N.  SYN. 

Subgenus  Hoplepicharis  Moure,  1945a.  Type-species:  E.fas- 
ciata  Lepeletier  and  Serville,  1828. 

74.  lunulata  Mocsary,  1899.  Mexico  to  Panama. 

75.  monozona  Mocsary,  1899.  Panama. 

Subgenus  Parepicharis  Moure,  1945a.  Type-species:  E.  zo- 
nata  F.  Smith,  1854. 

76.  metatarsalis  Friese,  1899.  Costa  Rica  to  Venezuela. 

= phenacura  Cockerell,  1917.  N.  SYN. 

= conura  Cockerell,  1917.  N.  SYN. 

Subgenus  Epicharoides  Radoszkowski,  1884.  Type-species: 
E.  maculata  F.  Smith,  1874. 

77.  albofasciata  F.  Smith,  1874.  Costa  Rica  to  South  Amer- 

ica. 

= nigroclypeata  Friese,  1899. 

78.  maculata  F.  Smith,  1874.  Mexico  to  northern  South 

America. 

= bipunctatus  Radoszkowski,  1884. 

= variabilis  Friese,  1900b. 

Subgenus  Epicharitides  Moure,  1945a.  Type-species:  E. 
cockerelli  Friese,  1900a. 

79.  undetermined  species  near  obscura  Friese.  Panama. 

ACKNOWLEDGMENTS 

For  the  use  of  material  cited  in  this  study  1 am  indebted  to: 
P.H.  Arnaud,  Jr.,  and  W.J.  Pulawski  (CAS),  R.W.  Brooks 
(RWB),  R.E.  Coville,  G.R.  Frankie,  E.G.  Linsley,  and  J.A. 
Powell  (UCB),  G.C.  Eickwort  (CORN),  H.E.  Evans  (MCZ), 
M.  Favreau  and  J.G.  Rozen,  Jr.  (AMNH),  S.S.  Gingras  and 
the  late  P.D.  Flurd,  Jr.  (USNM),  C.D.  Michener  (UKAN), 
A.R.  Moldenke  (ORSU),  J.L.  Neff  (NEFF),  D.  Roubik 
(ROUB),  R.O.  Schuster,  L.S.  Kimsey,  and  K.E.  Steiner  (UCD), 
L.A.  Stange  (DPIF),  and  T.J.  Zavortink  (TJZ). 

In  particular,  I wish  to  express  my  gratitude  to  G.R.  Else, 
S.S.  Gingras,  and  S.  Kelner-Pillault  for  making  available  crit- 
ical type  material  from  the  British  Museum  (Natural  His- 
tory), U.S.  National  Museum  of  Natural  History,  and  the 
Museum  d’Histoire  Naturelle,  respectively.  Special  thanks 
also,  to  R.W.  Brooks  for  important  notes  on  Friese  types  at 
the  Berlin  Museum. 

For  critically  reading  the  manuscript  and  offering  helpful 
suggestions  I am  indebted  to  C.L.  Hogue  and  to  the  various 
reviewers. 

Finally,  I wish  to  express  my  thanks  to  Beatriz  Larrain  for 
providing  the  Spanish  Resumen  for  this  paper. 

LITERATURE  CITED 

Alcock,  J.,  C.E.  Jones,  and  S.L.  Buchmann.  1976.  Location 
before  emergence  of  the  female  bee,  Centris  pallida,  by 
its  male  (Hymenoptera:  Anthophoridae).  Journal  of  Zo- 
ology 179:189-199. 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  67 


. 1977.  Male  mating  strategies  in  the  bee  Centris 

pallida  Fox  (Anthophoridae:  Hymenoptera).  The  Amer- 
ican Naturalist  1 1 1:145-155. 

Cameron,  P.  1903.  Descriptions  of  new  species  of  Hyme- 
noptera taken  by  Mr.  Edward  Whymper  on  the  “Higher 
Andes  of  the  Equator.”  Transactions,  American  Ento- 
mological Society,  29:225-238. 

Cheesman,  L.E.  1929.  Hymenoptera  collected  on  the  “St. 
George”  expedition  in  Central  America  and  the  W.  In- 
dies. Transactions,  Entomological  Society  of  London  77: 
141-154. 

Cockerell,  T.D.  A.  1899.  Note  on  the  nomenclature  of  some 
Hymenoptera.  The  Entomologist  32:14. 

. 1906.  The  North  American  bees  of  the  family  An- 
thophoridae. Transactions,  American  Entomological 
Society  32:63-1 16. 

. 1912.  New  bees  from  Brazil.  Psyche  19:41-61. 

. 1917.  New  bees  from  Costa  Rica  (Hym.).  Ento- 
mological News  28:200-201. 

. 1919.  Bees  in  the  collection  of  the  United  States 

National  Museum.  — 3.  Proceedings,  United  States  Na- 
tional Museum  55:167-221. 

. 1923.  Expedition  of  the  California  Academy  of 

Sciences  to  the  Gulf  of  California  in  1921.  The  Bees  (I). 
Proceedings,  California  Academy  of  Sciences  (ser.  4)  12: 
73-103. 

. 1924.  Anthophorid  bees  in  the  collection  of  the 

California  Academy  of  Sciences.  Pan-Pacific  Entomol- 
ogist 1:49-56. 

. 1949.  Bees  from  Central  America,  principally  Hon- 
duras. Proceedings,  United  States  National  Museum  98: 
429-490. 

Coville,  R.E.,  G.W.  Frankie,  and  S.B.  Vinson.  1983.  Nests 
of  Centris  segregata  (Hymenoptera:  Anthophoridae)  with 
a review  of  the  nesting  habits  of  the  genus.  Journal, 
Kansas  Entomological  Society  56:109-122. 

Crawford,  J.C.  1906.  Some  Costa  Rican  bees.  Transactions, 
American  Entomological  Society  32:157-163. 

. 1907.  Preoccupied  names  of  bees.  Canadian  Ento- 
mologist 39:21. 

Cresson,  E.T.  1865.  On  the  Hymenoptera  of  Cuba.  Pro- 
ceedings, Entomological  Society  of  Philadelphia  4: 1-200. 

. 1872.  Hymenoptera  Texana.  Transactions,  Amer- 
ican Entomological  Society  4:153-292. 

. 1879.  Descriptions  of  new  North  American  Hy- 
menoptera in  the  collection  of  the  American  Entomo- 
logical Society.  Transactions,  American  Entomological 
Society  7:201-214. 

Ducke,  A.  1902.  Beobachtungen  fiber  Blutenbesuch,  Er- 
scheinungszeit  etc.  der  bei  Para  vorkommenden  Bienen. 
Allegemeine  Zeitschrift  ffir  Entomologie  7:321-325, 
360-367,  400-404,  417-421. 

. 1907.  Contribution  a la  connaissance  de  la  fauna 

hymenopterologique  du  nord-est  du  Bresil.  Revue  d’En- 
tomologie,  Caen  26:73-76. 

Fabricius,  I.C.  1775.  Systema  entomologie.  Flensburg  and 
Leipzig,  xxx  + 832  pp. 


. 1793.  Entomologia  systematica  emendata  et  aucta. 

Copenhagen,  viii  + 519  pp. 

. 1804.  Systema  Piezatorum.  Braunschweig,  439  + 

30  pp. 

Fox,  W.J.  1899.  Synopsis  of  the  United  States  species  of 
the  hymenopterous  genus  Centris  Fabr.  with  description 
of  a new  species  from  Trinidad.  Proceedings,  Academy 
of  Natural  Sciences  of  Philadelphia  51:63-70. 

Friese,  H.  1899.  Neue  Arten  der  Bienengattungen  Epicharis 
Klug  und  Centris  Fabr.  Termeszetrajzi  FUzetek  22:39- 
48. 

. 1900a.  Neue  Arten  der  Bienengattungen  Centris 

und  Epicharis.  Termeszetrajzi  FUzetek  23:1 17-122. 

. 1900b.  Monographic  der  Bienengattung  Centris  (s. 

lat.).  Annalen,  K.K.  Naturhistorische  Hofmuseums,  Wien 
14:237-350. 

. 1904.  Nachtrag  zur  Monographie  du  Bienengat- 
tung Centris.  Annales  Musei  nationalis  Hungarici  2: 
90-92. 

. 1912.  Neue  und  wenig  bekannte  Bienenarten  der 

neotropischen  Region.  Archiv  ffir  Naturgeschichte  78(6): 
198-226. 

. 1913.  Uber  einige  neue  Apiden.  Archiv  Ffir  Na- 
turgeschichte 78(  1 2):85— 89. 

. 1925.  Neue  neotropische  Bienenarten,  zuglich  II. 

Nachtrag  zur  Bienenfauna  von  Costa  Rica  (Hym.).  Stet- 
tiner  Entomologische  Zeitung  86:1-41. 

Gnbodo,  G.  1894.  Note  imenotterologiche.  Nota  II,  con- 
tinuazione.  Bolletino,  Societa  Entomologica  Italiana  26: 
262-312. 

Harris,  R.A.  1979.  A glossary  of  surface  sculpturing.  Oc- 
casional Papers  in  Entomology,  California  Department 
of  Food  and  Agriculture,  28,  31  pp. 

Hurd,  P.D. , Jr.  1979.  In  K.V.  Krombein  et  al.  Catalog  of 
Hymenoptera  in  America  north  of  Mexico.  Smithsonian 
Institution,  Washington  2:xvi  + 2209  pp. 

International  Commission  of  Zoological  Nomenclature. 
1959.  Opinion  567.  (Centris  placed  on  Official  List  of 
Generic  Names  in  Zoology.) 

Klug,  J.C.F.  1807.  Vergleichung  der  Gattungen  der  Haut- 
flugler  Piezata  Fabr.  Hymenoptera  Linn.  Jur.  Magazin 
ffir  Insektenkunde  6:189-199. 

. 1810.  Einige  neue  Piezatengattungen.  Magazin  Ge- 

sellschaft  Naturforschender  Freunde  zu  Berlin  4:31-45. 

Latreille,  P.A.  1810.  Considerations  generates  . . . des  in- 
sectes.  Paris,  444  pp. 

Lepeletier  de  Saint-Fargeau,  A.  1841.  Histoire  naturelle  des 
insectes,  hymenopteres.  Paris,  2:1-680. 

Lepeletier  de  Saint-Fargeau,  A.,  and  A.  Serville.  1842.  In 
P.  A.  Latreille,  Encyclopedic  methodique.  Histoire  na- 
turelle, Entomologie.  Paris. 

Lutz,  F.E.,  and  T.D. A.  Cockerell.  1920.  Notes  on  the  dis- 
tribution and  bibliography  of  North  America  bees  of  the 
families  Apidae,  Meliponidae,  Bombidae,  Englossidae 
and  Anthophoridae.  Bulletin,  American  Museum  of 
Natural  History  42:491-641. 

Michener,  C.D.  1944.  Comparative  external  morphology, 


68  Contributions  in  Science,  Number  347 


Snelling:  American  Centridini 


phylogeny  and  a classification  of  the  bees  (Hymenop- 
tera).  Bulletin,  American  Museum  of  Natural  History 
82:151-326. 

. 1951.  Subgeneric  groups  of  Hemisia  (Hymenop- 

tera,  Apoidea).  Journal,  Kansas  Entomological  Society 
24:1-1 1. 

. 1954.  Bees  of  Panama.  Bulletin,  American  Mu- 
seum of  Natural  History  104:1-176. 

. 1965.  A classification  of  the  bees  of  the  Australian 

and  South  Pacific  Regions.  Bulletin,  American  Museum 
of  Natural  History  130:1-362. 

Mitchell,  T.B.  1962.  Bees  of  the  eastern  United  States. 
Technical  Bulletin,  North  Carolina  Agricultural  Exper- 
iment Station,  152,  557  pp. 

Mocsary,  A.  1898.  Species  novae  generum:  Eug/ossa  Latr. 
et  Epicharis  Klug.  Termeszetrajzi  Fiizetek  21:497-500. 

. 1899.  Species  novae  generis  Centris  Fabr.  Ter- 
meszetrajzi Fiizetek  22:251-255. 

Moure,  J.S.  1943.  Abelhas  de  Batatais  (Hym.,  Apoidea). 
Arquivas,  Museo  do  paranaense,  Curitiba  3:145-203. 

. 1945a.  Notas  sobre  as  Epicharitina  (Hymenop., 

Apoidea).  Revista  de  Entomologia  16:293-314. 

. 1945b.  Apoidea  da  Cole^ao  de  Conde  Amadeu  A. 

Barbiellini.  II  (Hym.  Apoidea).  Rcvista  de  Entomologia, 
16:394-414. 

. 1950.  Alguns  agrupamentos  novos  de  abelhas  Neo- 

tropicais.  Dusenia  1:385-394. 

. 1960a.  Abelhas  da  Regiao  Neotropical  descritas 

por  G.  Gribodo  (Hymenoptera-Apoidea).  Boletim,  Uni- 
versidade  do  Parana  1:1-18. 

. 1960b.  Notes  on  the  types  of  the  Neotropical  bees 

described  by  Fabricius  (Hymenoptera:  Apoidea).  Studia 
Entomologia  3:97-160. 

Moure,  J.S.,  and  C.A.C.  Seabra.  1959.  Notas  sobre  abelhas 
do  genero  Epicharis  (Hym.,  Apoidea).  Studia  Entomo- 
logia 2:1  19-127. 

Neff,  J.L.,  and  B.B.  Simpson.  1981.  Oil-collecting  struc- 
tures in  the  Anthophoridae  (Hymenoptera):  morphol- 
ogy, function,  and  use  in  systematics.  Journal,  Kansas 
Entomological  Society  54:95-123. 

Olivier,  G. A.  1789.  Encyclopedic  Methodique,  histoire  na- 
turelle  . . . Insectes.  Paris  and  Tiege,  4:cclxxxviij  + 
ccclxxiij  + 331  pp. 


Perez,  J.  1905.  Hymenopteres  nouveaux  du  Mexique.  Bul- 
letin, Museum  d'Histoire  Naturelle,  Paris  1 1:39-41. 

Rados/kowski,  O.  1884.  Quelques  nouveaux  Hymenop- 
teres d'Amerique.  Horae  Societe  Entomologique  Ros- 
sicae  18:17-22. 

Sandhouse,  G.A.  1943.  The  type  species  of  the  genera  and 
subgenera  of  bees.  Proceedings,  United  States  National 
Museum  92:519-619. 

Schwarz,  H.F.  1934.  The  solitary  bees  of  Barro  Colorado 
Island,  Canal  Zone.  American  Museum  Novitates  722: 
1-24. 

Smith,  F.  1854.  Catalogue  of  Hymenopterous  insects  in  the 
collection  of  the  British  Museum.  II,  Apidae.  British 
Museum.  London.  465  pp. 

. 1861.  Descriptions  of  new  genera  and  species  of 

exotic  Hymenoptera.  Journal  of  Entomology  1 : 1 46-155. 

. 1874.  A revision  of  the  genera  Epicharis,  Centris, 

Eulaema,  and  Eug/ossa  belonging  to  the  family  Apidae, 
section  Scopulipedes.  Annals  and  Magazine  of  Natural 
History  (ser.  4)  13:357-373. 

Snelling,  R.R.  1956.  Bees  of  the  genus  Centra  in  California 
(Hymenoptera:  Anthophoridae).  Pan-Pacific  Entomol- 
ogist 32: 1-8. 

. 1966.  The  taxonomy  and  nomenclature  of  some 

North  American  bees  of  the  genus  Centris  with  descrip- 
tion of  new  species  (Hymenoptera:  Anthophoridae).  Los 
Angeles  County  Museum  Contributions  in  Science  1 12: 
1-33. 

. 1974.  Notes  on  the  distribution  and  taxonomy  of 

some  North  American  Centris  (Hymenoptera:  Anthoph- 
oridae). Contributions  in  Science,  Los  Angeles  County 
Museum  of  Natural  History  259:1-41. 

. 1 980.  New  bees  of  the  genus  Hylaeus  from  Sri 

Lanka  and  India  (Hymenoptera:  Colletidae).  Contri- 
butions in  Science,  Los  Angeles  County  Museum  of  Nat- 
ural History  328:1-18. 

Timberlake,  P.H.  1940.  The  genus  Centris  in  California 
(Hymenoptera,  Apoidea).  Pan-Pacific  Entomologist  16: 
138-141. 


Submitted  18  April  1983;  accepted  31  August  1983. 


Contributions  in  Science,  Number  347 


Snelling:  American  Centridini  69 


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SALAMANDERS  OF  THE  NORTHWESTERN 
HIGHLANDS  OF  GUATEMALA 

Paul  Elias' 


ABSTRACT.  The  salamander  faunas  of  the  Cordillera  de  los  Cu- 
chumatanes  and  the  Montanas  de  Cuilco  of  western  Guatemala  are 
reviewed  in  light  of  extensive  new  collections.  Thirteen  species  are 
reported  from  the  Cuchumatanes  including  one  new  member  of  the 
Bolitoglossa  mexicana  species  group,  herein  described  as  Bolito- 
glossa  jacksoni  sp.  nov.,  and  four  recognized  species  never  before 
collected  in  the  massif  ( Bolitoglossa  hartwegi,  B.  mulleri,  B.  rufescens 
and  Dendrotriton  rabbi).  Four  species  are  reported  from  the  Cuilco 
including  two  hitherto  unknown  from  that  mountain  range  ( Boli- 
toglossa mono  and  Pseudoeurycea  rex). 

Bolitoglossa  resplendens  (sensu  lato)  lacks  the  features  purported 
to  differentiate  it  from  B.  lincolni  and  is  considered  a junior  syn- 
onym. Based  on  gross  morphological  similarity,  the  Bolitoglossa 
helmrichi  and  B.  rostrata  species  groups  are  divided  and  reconsti- 
tuted into  four  species  groups  (dunni, franklini.  morio  and  veracrucis 
groups). 

The  Cuchumatanes  support  fewer  salamander  species  but  a larger 
number  of  species  groups  and  genera  than  do  the  Pacific  Uplands. 
The  diversity  differences  are  correlated  with  greater  age  and  con- 
nectedness among  high  elevation  salamander  habitats  in  the  Cu- 
chumatanes relative  to  the  Pacific  Uplands. 

Salamander  species  on  the  Cuchumatanes  and  Cuilco  occur  in 
narrow  elevational  belts  as  they  do  on  the  Pacific  Uplands.  Con- 
specific  populations  or  related  species  tend  to  occur  at  comparable 
elevations  wherever  found.  Within-group  variation  in  elevational 
range  does  exist,  however,  and  much  of  it  is  correlated  with  geo- 
graphic variation  in  the  elevation  of  the  cloud  line.  The  Bolitoglossa 
veracrucis  species  group  is  exceptional  among  Nuclear  Central  Amer- 
ican salamander  groups  in  the  extreme  breadth  of  the  combined 
elevational  ranges  of  its  three  member  species  (2,800  m;  the  next 
broadest  is  1,200  m).  The  great  ecological  scope  of  the  Bolitoglossa 
veracrucis  group  may  have  been  a crucial  factor  in  the  invasion  of 
the  lowlands  by  Bolitoglossa  beta. 

RESUMEN.  Las  faunas  de  salamandras  de  la  Cordillera  de  los  Cu- 
chumatanes y de  las  Montanas  de  Cuilco  en  el  oeste  de  Guatemala 
se  revisan  en  base  a las  extensas  nuevas  colecciones  realizadas.  Trece 
especies  se  encuentran  en  los  Cuchumatanes,  incluyendo  una  nueva 
especie,  Bolitoglossa  jacksoni,  miembro  del  grupo  espedfico  de  Bo- 
litoglossa mexicana,  y cuatro  especies  ya  descritas  pero  nunca  en- 
contradas  en  esta  sierra  previamente  (Bolitoglossa  hartweig,  B. 

1.  Museum  of  Vertebrate  Zoology,  University  of  California, 
Berkeley,  California  94720. 


mulleri,  B rufescens  y Dendrotriton  rabbi).  Cuatro  especies  se 
encuentran  en  el  Cuilco  incluyendo  dos  anteriormente  desconocidas 
en  esta  region  (Bolitoglossa  morio  y Pseudoeurycea  rex). 

Bolitoglossa  resplendens  (sensu  lato)  no  tiene  las  caracteristicas 
que  supuestamente  la  diferencian  de  la  B.  lincolni,  y consecuente- 
mente  B.  resplendens  es  un  sinonimo.  En  base  de  similaridades  en 
morfologia  externa,  los  grupos  especificos  de  Bolitoglossa  helmrichi 
y B.  rostrata  descritos  por  Wake  y Lynch  (1976)  son  divididos  y 
redefinidos  en  cuatro  grupos  (dunni,  franklini,  morio  y veracrucis). 

En  los  Cuchumatanes  habitan  menos  especies  de  salamandras  pero 
un  numero  mayor  de  grupos  especificos  y generos,  en  comparacion 
con  las  montafias  del  lado  Pacifico.  Estas  diferencias  de  diversidad 
estan  correlacionadas  con  una  edad  geologica  mayor  y con  una  mayor 
conexion  entre  regiones  de  alta  elevacion  en  comparacion  con  las 
montafias  del  lado  Pacifico. 

Las  especies  de  urodelos  en  los  Cuchumatanes  y el  Cuilco  se  lo- 
calizan  en  estrechas  bandas  elevacionales  al  igual  que  en  las  mon- 
tafias del  Pacifico.  Poblaciones  de  la  misma  especie,  o especies  re- 
lacionadas,  generalmente  se  encuentran  en  la  misma  zona  elevacional. 
Pero,  si  se  encuentra  variacion  al  nivel  de  intra-grupo,  la  mayor 
parte  de  esta  variacion  esta  correlacionada  con  la  variacion  geografica 
a la  altura  de  la  zona  nublada.  El  grupo  espedfico  de  Bolitoglossa 
veracrucis  es  una  exception  entre  todos  los  grupos  de  Centro  America 
Nuclear  debido  a la  extrema  distribucion  elevacional  de  las  tres 
especies  miembros  (2,800  m);  la  segunda  distribucion  mas  amplia 
es  1,200  m.  El  gran  alcance  geologico  de  este  grupo  puede  tener  un 
papel  importante  en  la  historia  de  la  invasion  de  las  tierras  bajas 
por  Bolitoglossa  beta. 

INTRODUCTION 

Nuclear  Central  America  (NCA),  the  landmass  bounded  to 
the  northwest  by  the  Isthmus  of  Tehuantepec  and  to  the 
southeast  by  the  Nicaraguan  depression,  contains  six  major 
highland  masses  isolated  from  one  another  by  deep  dry  val- 
leys (Fig.  1).  The  Cordillera  de  los  Cuchumatanes  and  the 
Montanas  de  Cuilco  are  two  limestone  capped  ranges,  re- 
spectively about  3,000  and  400  square  kilometers  in  extent 
and  4,000  and  3,400  meters  at  greatest  elevation  (Fig.  2). 

On  the  Caribbean  slope  of  the  Cuchumatanes,  tropical  rain 
forest  (following  the  ecological  terminology  of  Schmidt,  1 936) 
extends  from  near  sea  level  to  a 1,300  m "cloud  line”  (that 
characteristic  elevation  at  which  the  water  carried  by  rising 


Contributions  in  Science,  Number  348,  pp.  1-20 
Natural  History  Museum  of  Los  Angeles  County,  1984 


ISSN  0459-8113 


humid  air  condenses  in  the  form  of  fog).  Above  the  cloud 
line  is  a zone  of  extremely  wet  broadleaf  and  treefern  forest, 
the  subtropical  zone  cloud  forest,  which  grades  upward  into 
drier  but  still  humid  subtropical  forest  that  is  frequently  com- 
posed of  oak.  At  about  2,700  m,  varying  locally,  the  pine 
and  cypress  temperate  zone  forests  commence,  continuing 
to  the  3,400-m  treeline,  above  which  bunchgrass  predomi- 
nates. 

The  above  vegetational  series  is  found  on  the  unobstructed 
face  of  the  Caribbean  slope;  but,  in  the  protected  interior 
valleys,  variably  drier  systems  occur,  typically  dominated  by 
pine-oak  forest  even  at  low  elevation.  Because  of  their  pro- 
tected position,  the  Montanas  de  Cuilco  have  an  abbreviated 
form  of  the  ecological  pattern  found  on  the  open  escarpment: 
as  in  the  Cuchumatanes,  pine  and  cypress  dominate  down 
to  about  2,700  m,  where  a wet  broadleaf  forest  begins;  how- 
ever, a high  cloud  line  occurs  at  about  2,000  m,  and  dry 
scrubby  pine-oak  growth  predominates  below  this  level. 

The  salamander  faunas  of  the  Cordillera  de  los  Cuchu- 
matanes and  the  Montanas  de  Cuilco  of  western  Guatemala 
have  been  little  sampled  or  reported  upon.  In  the  Cuchu- 
matanes Stuart  ( 1 943a,  1 943b)  made  the  first  collections  and 
reported  four  species  referred  to  Oedipus : lincolni,  cuchu- 
matanus,  rostratus,  and  rex.  From  later  collections  Stuart 
(1952)  described  Magnadigita  omniumsanctorum.  The  next 
figure  1.  Nuclear  Cential  America.  The  2,100-m  contour  interval  discovery  in  the  Cuchumatanes  was  recorded  by  Lynch  and 
is  suppled. 


Figure  2.  Montanas  de  Cuilco  and  the  Cordillera  de  los  Cuchumatanes  with  the  2,100-m  contour  interval  and  regional  landmarks. 


2 Contributions  in  Science,  Number  348 


Elias:  Guatemalan  Salamanders 


Figure  3.  Bolitoglossa  rostrata,  B.  cuchumatana,  B.  mono,  and  B.  lincolni,  life  size.  UPPER  LEFT:  Boliloglossa  rostrata,  body,  legs,  and 
flanks  dark  brown,  bilateral  shoulder  stripes  lighter  brown  with  middorsal  brown  mottling.  A unicolor  brown  morph  also  is  found.  UPPER 
LEFT  INSET:  Feet  of  B.  cuchumatana.  Overall  coloration  like  that  of  B.  rostrata.  UPPER  RIGHT:  B.  morio,  dark  slate  grey  all  over  except 
for  light  speckling  on  flanks  and  anterior  surfaces  of  limbs,  and  light  lateral  spotting  on  tail  (Cuilco  specimen  figured;  Cuchumatan  specimens 
have  light  spotting  on  flanks  as  well  as  tail).  BOTTOM  CENTER:  B lincolni,  black  ground  color  with  a coral  red  dorsal  swath  and  spotting 
on  limbs  and  (occasionally)  venter.  In  some  specimens,  the  dorsal  swath  is  reduced  to  a series  of  red  spots. 


Wake  (1975)  with  the  naming  of  Chiropterotriton  cuchu- 
matanus.  Finally,  Elias  and  Wake  (1983)  and  Wake  and  Elias 
(1983)  named  two  new  monotypic  genera,  Nyctanolis  pernix 
and  Bradytriton  silus,  both  from  the  cordillera. 

Prior  to  the  present  study  the  only  salamanders  that  had 


been  collected  in  the  Montanas  de  Cuilco  were  Bolitoglossa 
resplendens  ( sensu  lato)  and  Chiropterotriton  rabbi,  both  first 
reported  by  Lynch  and  Wake  (1975). 

Bolitoglossa  stuarti  was  described  from  the  valley  sepa- 
rating the  Cuilco  from  the  Cuchumatanes  (Wake  and  Brame, 


Contributions  in  Science,  Number  348 


Elias:  Guatemalan  Salamanders  3 


□ 


Figure  4.  Relationship  between  maxillary  tooth  number  (both  sides 
summed)  and  standard  length  in  the  Bolitog/ossa  dunni  complex. 
Solid  triangle  = B.  cuchumatana  (Eastern  Cuchumatanes);  open  tri- 
angle = B.  cuchumatana  (Western  Cuchumatanes);  half-filled  cir- 
cle = B.  helmrichi  (Chuacus);  solid  circle  = B helmrichi  (Western 
Minas);  open  circle  = B.  helmrichi  (Xucaneb);  solid  square  = B.  en- 
gelhardti  (Pacific  Uplands);  open  square  = B.  sp.  (Eastern  Cuchu- 
matanes); half-filled  square  = B.  dunni  (Elonduras). 

1969)  and  is  the  only  salamander  species  known  front  the 
valley.  As  a low  elevation  form  this  species  will  be  mentioned 
only  in  the  discussion. 

In  the  course  of  the  present  study  twelve  salamander  species 
were  documented  for  the  Cuchumatanes,  bringing  the  total 
number  of  known  forms  from  six  to  thirteen  for  the  mountain 
range.  Among  the  seven  species  previously  unknown  from 
the  massif  were  the  two  new  monotypic  genera  described 
elsewhere  (Elias  and  Wake,  1983;  Wake  and  Elias,  1983)  and 
the  one  new  member  of  the  Bo/itoglossa  mexicana  species 
group  described  below.  Four  species  were  taken  in  the  Mon- 
tanas de  Cuilco  during  this  work.  All  of  these  were  recognized 
forms,  but  two  were  previously  unknown  from  those  moun- 
tains. 

This  paper  provides  a species  account  for  each  of  the  sal- 
amander species  now  known  from  the  Cuchumatanes  and 
Cuilco,  including  taxonomic  synonymies,  type  locality,  geo- 
graphic range,  habits  and  habitat  preferences  and  a discussion 
of  the  geographic  variation  and  possible  evolutionary  rela- 
tionships of  the  taxon.  The  discussion  section  is  divided  into 
three  parts.  In  the  first  part,  the  species  groups  within  Boli- 
toglossa  beta  ( sensu  Wake  and  Lynch,  1976)  are  redefined. 
In  the  second  part,  contemporary  patterns  of  distribution 
and  differentiation  are  examined  to  provide  insight  into  the 
history  of  the  salamander  fauna.  In  the  third  part,  patterns 
of  elevational  distribution  are  presented  for  ihe  Cuchuma- 


tanes and  Cuilco  and  are  discussed  relative  to  the  results  of 
Wake  and  Lynch  (1976)  for  the  Pacific  Uplands. 

SPECIES  ACCOUNTS 

Bolitoglossa  cuchumatana  (Stuart,  1943) 

Figure  3 

Oedipus  cuchumatanus  Stuart,  1943a:  14. 

Magnadigita  cuchumatana:  Taylor,  1944:218. 

Bolitoglossa  cuchumatana:  Wake  and  Brame,  1963:386. 

TYPE  LOCALITY.  Oak  woods  2 km  N Nebaj,  Depto. 
Quiche,  Guatemala,  1,900  m elevation. 

RANGE.  Stuart  collected  this  form  only  at  the  type  lo- 
cality. Wake,  Houck,  and  Lynch  next  found  the  species  25 
years  later  about  15  km  to  the  south  and  600  m higher  in 
elevation.  I found  B.  cuchumatana  at  two  additional  sites  in 
the  western  Cuchumatanes  near  Chiapas  and  at  the  eastern- 
most end  of  the  range,  near  San  Miguel  Uspantan. 

HABITAT  AND  HABITS.  B.  cuchumatana  have  been 
taken  on  the  ground,  under  bark,  and  in  arboreal  bromeliads 
where  they  have  been  found  active  on  surface  debris  on  wet 
evenings  from  the  1 ,200-m  cloud  line  through  the  subtropical 
cloud  forest  to  2,500  m. 

SYSTEMATICS.  A series  of  species,  including  Bolito- 
glossa engelhardti  in  the  Pacific  Uplands,  B.  helmrichi  in  the 
Xucaneb  and  Western  Minas,  a newly  discovered  population 
(possibly  representing  a new  species)  in  the  Eastern  Minas, 
B.  cuchumatana  in  the  Cuchumatanes,  and  B.  dunni  in  Hon- 


Table  1.  Geographic  variation  in  subcaudal  coloration  and  degree 
of  interdigital  webbing  among  the  species  of  the  Boltioglossa  dunni 
complex. 


Species  and 
population 

Subcaudal 

coloration 

Degree  of 
interdigital 
webbing 

B.  helmrichi 
(Chuacus) 

no  data 

high 

B.  helmrichi 
(Xucaneb) 

orange 

high 

B.  helmrichi 
(Western  Minas) 

orange 

high 

B.  sp. 

(Eastern  Minas) 

tan 

low 

B.  cuchumatana 
(Western  Cuchumatanes) 

tan 

medium 

B.  cuchumatana 
(Eastern  Cuchumatanes) 

tan 

medium 

B.  engelhardti 
(Pacific  Uplands) 

tan 

medium 

B.  dunni 
(Honduras) 

no  data 

medium 

4 Contributions  in  Science,  Number  348 


Elias:  Guatemalan  Salamanders 


Figure  5.  BolHoglossa  jacksoni,  B.  rufescens,  B.  mulleri,  Dendrotriton  rabbi,  D.  cuchumatanus,  and  Boliloglossa  hartwegi.  life  size.  UPPER 
LEFT:  Boliloglossa  jacksoni  (holotype  figured),  background  yolky  yellow,  middorsal  swath  dark  brown  with  narrow  white  border.  TOP 
CENTER:  B.  rufescens.  tan  overall  with  a slightly  darker  brown  snout  and  flanks.  Tone  varies  front  light  to  dark  brown  both  between  individuals 
and  for  one  individual  at  different  times  of  day.  UPPER  RIGHT:  B.  mulleri,  black  overall  except  for  a narrow  broken  vertebral  stripe  of  tan. 
In  some  Cuchumatan  specimens,  the  stripe  is  reduced  or  absent.  LOWER  LEFT:  Dendrotriton  rabbi,  pattern  composed  of  various  shades  of 
brown.  Note  light  interorbital  bar.  Animals  may  have  a reddish  or  even  greenish  cast.  Pattern  and  tone  variable  between  individuals.  BOTTOM 
CENTER:  D.  cuchumatanus,  similar  to  D.  rabbi  above.  LOWER  RIGHT:  Bo/itoglossa  hartwegi,  dark  brown  overall  with  lighter  paired 
shoulder  and  pelvic  stripes.  Animals  may  be  without  markings  altogether  or  may  have  small  light  streaks  on  dorsum  at  random. 


Contributions  in  Science,  Number  348 


Elias:  Guatemalan  Salamanders  5 


Figure  6.  Bolitoglossa  jacksoni  sp.  nov.,  an  adult  female  (escaped)  from  the  type  locality. 


duras,  appears  to  form  a relatively  homogeneous  assemblage 
of  related  congeners.  The  group,  which  I shall  call  the  dunni 
complex,  was  first  outlined  by  Stuart  (1943a)  and  is  char- 
acterized by  a moderate  degree  of  foot  webbing;  by  well- 
developed  subdigital  pads  and  the  associated  hypertrophy  of 
the  terminal  phalanges;  by  a generally  brown  dorsal  color 
frequently  differentiated  into  light  shoulder  stripes  and  dark- 
er flanks;  and  by  a common  microhabitat  (see  Wake  and 
Lynch,  1976,  for  information  on  the  ecology  of  B.  engel- 
hardti). 

The  species  in  this  series  are  most  similar  to  Bolitoglossa 
rostrata,  but  differences  in  foot  morphology  (B.  rostrata  has 
little  mterdigital  webbing)  usually  permit  separation  (see  B. 
rostrata  and  compare  inset  B.  cuchumatana  feet.  Fig.  3).  B. 
rostrata  occurs  in  limited  sympatry  with  B.  cuchumatana 
with  no  hybridization. 

Despite  the  general  similarities,  variation  between  the 
species  of  the  B.  dunni  complex  is  pronounced  in  the  col- 
oration of  the  subcaudal  surface,  the  number  of  maxillary 
teeth,  and  the  morphology  of  the  foot.  Data  were  collected 
for  eight  populations  from  the  five  B.  dunni  complex  species 
(Fig.  4,  Table  1).  B.  helmrichi  can  be  defined  on  the  basis  of 
its  high  degree  of  interdigital  webbing  and  orange  subcaudal 
coloration  to  include  populations  in  the  Xucaneb,  the  West- 
ern Minas,  and  the  Chuacus.  B.  cuchumatana , including  all 
Cuchumatan  specimens,  and  B.  engelhardti  are  similar  in 
their  intermediate  degree  of  interdigital  webbing,  tan  sub- 
caudal surface,  and  moderate  tooth  number,  and  are  distin- 
guishable from  one  another  both  on  the  basis  of  adult  size 


(B.  engelhardti  is  smaller  than  any  other  member  of  the  com- 
plex) and  in  frequency  of  shoulder  striping  ( B . engelhardti  is 
usually  unmarked,  while  B.  cuchumatana  almost  always  has 
bilateral  light  striping).  Bolitoglossa  dunni  occurs  in  both  a 
pink  and  dark  morph,  the  latter  with  light  spotting  (Schmidt, 
1933),  and  it  achieves  larger  size  than  any  other  member  of 
the  complex  and  has  little  webbing  and  low  tooth  numbers. 
The  population  from  the  Eastern  Minas  differs  sharply  from 
all  other  members  of  the  B.  dunni  complex  (and  from  all 
known  B.  rostrata  populations  as  well)  in  having  extremely 
high  numbers  of  maxillary  teeth.  It  has  almost  as  little  web- 
bing as  B.  rostrata.  That  this  population  belongs  in  the  B. 
dunni  complex  rather  than  to  B.  rostrata  is  suggested  only 
by  its  occurrence  at  relatively  low  elevation. 

Bolitoglossa  hartwegi  Wake  and  Brame,  1969 

Figure  5 

Bolitoglossa  hartwegi  Wake  and  Brame,  1969:10. 

TYPE  LOCALITY.  4.5  mi.  W San  Cristobal  de  Las  Casas, 
Chiapas,  Mexico.  7,000  ft  elevation. 

RANGE.  B.  hartwegi  has  been  known  only  from  the  gen- 
eral vicinity  of  its  type  locality  on  the  Mesa  Central  de  Chia- 
pas. It  is  here  reported  from  Guatemala,  both  in  the  western 
Cuchumatanes  at  Finca  Chiblac,  and  in  the  easternmost 
reaches  of  the  range,  near  San  Miguel  Uspantan. 

HABITAT  AND  HABITS.  Specimens  have  been  collect- 
ed under  bark  on  logs  and  stumps,  under  cover  objects  on 
the  ground,  in  rock  crevices,  and  in  arboreal  bromeliads;  the 


6 Contributions  in  Science,  Number  348 


Elias:  Guatemalan  Salamanders 


species  is  active  on  rainy  nights.  B.  hartwegi  have  a broad 
elevational  range,  occurring  from  the  1,200-m  cloud  line  to 
the  lower  temperate  forest  at  over  2,700  m. 

SYSTEMATICS.  When  these  populations  were  discov- 
ered in  the  Cuchumatanes  during  this  study  they  were  as- 
signed to  the  Bolitoglossa  veracrucis  species  group  (see  Dis- 
cussion) based  upon  their  well-ossified  skull  with  a broad 
pars  dentalis  of  the  premaxillary,  and  a derived  foot  mor- 
phology. They  were  assigned  to  the  species  B.  hartwegi  based 
on  a mainly  unicolor  dorsum  ( B . veracrucis  is  streaked  and 
mottled),  a small  size  (B.  stuarti  is  larger),  and  a mottled 
venter  (unicolor  in  B.  stuarti). 

Bolitoglossa  jacksoni  new  species 

Figures  5 and  6 

HOLOTYPE.  MVZ  134634,  a young  adult  female  from 
the  Las  Nubes  sector  of  Finca  Chiblac,  approximately  1 2 km 
NNE  of  Santa  Cruz  Barillas,  Depto.  Fluehuetenango,  Gua- 
temala, at  about  1,400  m elevation,  collected  by  Jeremy  L. 
Jackson,  1 September  1975. 

DIAGNOSIS.  The  new  taxon  is  a large  species  of  Boli- 
toglossa (the  young  adult  holotype  is  49.1  mm  and  another 
specimen  was  approximately  65  mm  in  standard  length)  with 
low  numbers  of  maxillary  teeth  (31  in  the  holotype)  and 
moderate  numbers  of  vomerine  teeth  (21  in  the  holotype). 
The  species  is  a member  of  the  Bolitoglossa  alpha  group  on 
the  basis  of  its  simple  vertebral  tail  autotomy  mechanism 
(Wake  and  Dresner,  1967;  Wake  and  Lynch,  1976).  It  is 
distinguished  from  B.  salvinii,  B.  mexicana.  B.  mulleri,  B. 
odonne/li.  B.  platydactyla.  and  B.  flaviventris  in  having  dark 
coloration  restricted  to  the  middorsal  region;  separated  from 
all  other  Nuclear  Central  American  Bolitoglossa  both  by  col- 
or pattern  and  its  tail  autotomy  mechanism. 

DESCRIPTION  (OF  HOLOTYPE).  Nostril  small,  labial 
protuberances  of  nasolabial  grooves  small  and  poorly  de- 
veloped; canthus  rostralis  moderate  in  length,  rounded  and 
gently  arched.  Standard  length  6.6  times  head  width;  stan- 
dard length  4.5  times  snout-gular  fold  length  (head  length). 
Deep,  unpigmented  groove  below  eye  extends  almost  full 
length  of  opening,  following  curvature  of  eye,  but  does  not 
extend  to  lip.  Eye  moderate  in  size,  slightly  protuberant  in 
life.  Lightly  indicated  postorbital  groove  extends  posteriad 
and  slightly  ventrad  as  a shallow  depression  for  2.7  mm, 
thence  proceeds  directly  ventrad  and  passes  behind  posterior 
end  of  mandible  to  produce  a shallow  but  definite  nuchal 
groove  parallel  to  and  4.0  mm  anterior  to  the  dearly  defined 
gular  fold.  The  21  vomerine  teeth  are  arranged  in  single 
arched  rows  on  either  side  of  the  midline  and  extend  laterally 
to  the  level  of  the  midpoint  of  the  internal  nares.  The  two 
rows  of  3 1 maxillary  teeth  extend  posteriad  to  a level  one- 
third  of  the  way  through  the  eye.  Five  small  undifferentiated 
premaxillary  teeth  protrude  from  well  behind  the  lip.  The 
tail  is  round  in  cross  section  and  is  0.8  times  the  standard 
length  from  base  to  tip;  the  postiliac  gland  is  not  evident. 
Limbs  are  of  moderate  length,  three  costal  grooves  are  ex- 
posed between  the  toetips  of  the  adpressed  fore  and  hind 
limbs;  standard  length  is  4.4  times  right  hind  limb  length 


and  4.9  times  right  fore  limb  length.  Standard  length  is  10.0 
times  the  hind  foot  width  and  both  fore  and  hind  feet  are 
fully  webbed  and  lack  subdigital  pads;  the  toetips  are  rounded 
in  dorsal  aspect.  The  fingers  in  order  of  decreasing  length:  3, 
2,  4,  1;  toes  in  order  of  decreasing  length:  3,  4,  2,  5,  1. 

MEASUREMENTS  (OF  HOLOTYPE).  Head  width 
(maximum),  7.4  mm;  head  length  (snout  to  gular  fold),  1 1.0 
mm;  head  depth  at  posterior  angle  of  jaw,  4.1  mm;  eyelid 
length,  3.0  mm;  eyelid  width,  1.9  mm;  anterior  rim  of  orbit 
to  snout,  2.6  mm;  horizontal  orbital  diameter,  2.2  mm;  m- 
terorbital  distance,  3.0  mm;  distance  between  vomerine  teeth 
and  parasphenoid  tooth  patch,  0.3  mm;  snout  to  fore  limb, 

13.6  mm;  distance  separating  internal  nares,  2.2  mm;  dis- 
tance separating  external  nares,  2.1  mm;  snout  projection 
beyond  mandible,  0.6  mm;  snout  to  posterior  angle  of  vent 
(standard  length),  49.1  mm;  snout  to  anterior  angle  of  vent, 

45.6  mm;  axilla  to  groin,  27.4  mm;  tail  length,  43.4  mm;  tail 
width  at  base,  3.7  mm;  tail  depth  at  base,  3.3  mm;  fore  limb 
length,  10.0  mm;  hind  limb  length,  1 1.2  mm;  width  of  hand, 
3.7  mm;  width  of  foot,  4.9  mm. 

COLORATION  IN  ALCOHOL.  The  ground  color  is  pale 
yellow  with  a broad  middorsal  swath  of  dark  brown.  The 
brown  marking  originates  on  the  head  at  the  level  of  the 
eyelids  and  extends  in  breadth  from  the  center  of  one  eyelid 
to  the  center  of  the  other  in  a straight  edge  across  the  head. 
Posteriorly  the  brown  stripe  diminishes  gradually  until  it 
disappears  near  the  tail  tip.  Aside  from  the  brown  swath,  the 
animal  is  unmarked  (Fig.  5). 

COLORATION  IN  LIFE.  This  animal  was  a brilliant  yolky 
yellow.  The  dorsal  swath  was  chocolate  brown  and  was  en- 
tirely bordered  by  an  edging  of  immaculate  white  less  than 
a millimeter  in  width.  The  eye  was  metallic  gold  (Fig.  6). 

OSTEOLOGY.  As  the  animal  was  preserved  flat  in  stan- 
dard pose,  accurate  measurements  from  a radiograph  could 
be  taken.  Skull,  from  the  anterior  border  of  the  premaxilla 
to  the  medial  edge  of  the  occipital  condyles,  8.0  mm;  width 
of  pars  dentalis  of  the  premaxilla,  1.3  mm;  maximum  width 
of  braincase  (anterior  to  the  otic  capsules)  3.0  mm;  maximum 
width  of  skull  base  (across  otic  capsules)  4.9  mm;  longest 
axis  of  otic  capsule,  2.6  mm;  and  straight  line  length  of  the 
maxilla  from  anterior  to  posterior  tip,  4.5  mm. 

The  preorbital  process  of  the  vomer  extends  well  lateral 
to  the  internal  narial  opening  as  a slender  sliver  that  tapers 
to  a point.  The  frontal  processes  of  the  premaxilla  arise  and 
continue  well  separated  from  one  another,  spreading  grad- 
ually as  they  rise  dorsad  and  posteriad.  Septomaxillae  are 
absent.  Trunk  vertebrae  number  14,  caudosacral  2,  and  cau- 
dal 30.  No  tibial  spur  is  evident;  phalangeal  formulae  are  1, 
2,  3,  3,  2 for  the  foot  and  1,  2,  3,  2 for  the  hand.  Phalanges 
are  broadened  and  slightly  irregular  in  outline,  and  the  ter- 
minals of  the  three  outer  fingers  and  four  outer  toes  are 
laterally  expanded  such  that  each  is  broader  than  it  is  long. 
The  innermost  toe  and  finger  have  terminals  that  are  pointed 
and  curved  along  the  rim  of  the  pad  toward  the  other  digits. 

VARIATION.  Two  specimens  of  this  form  were  collected, 
but  one,  a large  adult  female,  escaped  before  it  could  be 
preserved.  Knowledge  of  variation  is  thus  restricted  to  as- 
pects of  color  and  pattern  that  were  preserved  in  photographs 


Contributions  in  Science,  Number  348 


Elias:  Guatemalan  Salamanders  7 


98°  96°  94°  92°  90°  88° 

Figure  7.  Distribution  of  the  Boliloglossa  mexicana  species  group  (modified  front  Wake  and  Lynch,  1976). 


of  the  escaped  animal  (Fig.  6,  and  Hanken  et  al.,  1980).  All 
the  hues  described  for  the  holotype  were  similar  in  the  other 
individual,  but  the  latter  had  a more  irregular  dorsal  pattern. 
The  brown  dorsal  swath  was  interrupted  by  a large  yellow 
nape  spot  and  a break  on  the  tail,  and  the  white  border  was 
incomplete.  In  addition  a spot  of  brown  was  present  on  the 
dorsal  surface  of  the  right  hind  foot. 

RANGE.  Bolitoglossa  jacksoni  has  been  taken  only  within 
1 km  of  the  type  locality  on  the  Caribbean  escarpment  of 
the  western  Cuchumatanes  (Fig.  7). 

HABITAT  AND  HABITS.  Both  individuals  were  taken 
under  bark  of  felled  hardwood  logs  in  clearings  in  a sub- 
tropical forest,  which  receives  upward  of  6 m of  rain  annually. 

SYSTEMATICS.  Bolitoglossa  jacksoni  is  a member  of  the 
B.  mexicana  species  group  ( sensu  Wake  and  Lynch,  1976) 
which  also  includes  B.  Jlaviventris,  B.  mexicana,  B.  mulleri, 
B.  odonnelli,  B.  platydactyla,  and  B.  salvinii.  The  members 
of  the  group  share  a variety  of  structural  and  ecological  traits; 
all  have  the  primitive  (alpha  type)  vertebral  tail  autotomy 
mechanism  (Wake  and  Dresner,  1 967);  all  have  fully  webbed 
feet  that  lack  subdigital  pads  but  have  expanded  terminal 
phalanges;  all  are  large  species,  each  represented  by  speci- 
mens exceeding  65  mm  standard  length;  and  they  occur  only 
in  the  tropical  and  low  subtropical  forests. 


The  species  of  the  B.  mexicana  group  are  similar  in  general 
structure  and  have  been  defined  mainly  on  the  basis  of  col- 
oration. The  color  patterns  of  the  six  species  are  described 
below;  their  geographic  ranges  are  indicated  in  Figure  7. 

Bolitoglossa  Jlaviventris  typically  has  paired  dorsolateral 
light  stripes  that  begin  on  each  eyelid  and  join  to  produce  a 
light  snout.  In  the  extreme  condition,  these  stripes  are  paired 
all  the  way  onto  the  tail,  but  often  they  are  irregularly  de- 
veloped and  may  form  a light  reticulation,  especially  pos- 
teriorly. This  species  differs  from  others  with  dark  flanks  in 
having  a light  colored  and  unmarked  ventral  surface.  The 
light  ventral  coloration  sometimes  invades  the  dorsal  sur- 
faces of  the  legs  as  well,  but  the  flanks  are  always  dark  and 
sharply  set  off  from  the  light  belly  (see  Wake  and  Lynch, 
1976,  fig.  24).  The  eye  is  golden. 

Bolitoglossa  mexicana  has  a broad  middorsal  light  swath 
that  is  typically  broken  into  three  longitudinal  stripes  by  the 
invasion  of  dark  ground  color  onto  the  dorsal  surface.  The 
dorsal  pattern  may  range  from  virtually  a single  swath  (as  in 
B.  platydactyla ) to  triple  stripes,  and  then,  through  loss  of 
the  vertebral  line,  to  a bilinear  pattern  much  as  seen  in  B. 
odonnelli  (described  below).  The  snout  is  heavily  flecked, 
and  light  flecking  extends  over  all  of  the  dark  regions  of  the 
body  (see  Wake  and  Lynch,  1976,  frontispiece;  Taylor  and 


8 Contributions  in  Science,  Number  348 


Elias:  Guatemalan  Salamanders 


Smith,  1945,  fig.  58;  Stuart,  1943a,  plate  II,  fig.  2).  The  eye 
is  dark. 

Bolitoglossa  mulleri  typically  has  a narrow  vertebral  light 
stripe.  When  maximally  developed,  the  stripe  extends  from 
the  tail  tip  to  the  nape  of  the  neck  where  it  bifurcates  before 
terminating  on  the  eyelids.  The  light  pigmentation  is  fre- 
quently reduced,  and  may  be  absent  altogether  leaving  a 
uniformly  black  animal  (see  Fig.  5 and  Stuart,  1943a,  plate 
II,  fig.  4).  The  eye  is  dark. 

Bolitoglossa  odonnelh  has  narrow  paired  light  stripes.  At 
their  maximum  development,  these  stripes  extend  continu- 
ously from  the  eyelids  to  the  tip  of  the  tail,  but  frequently 
they  are  broken  into  rows  of  dots  or  may  not  extend  as  far 
anteriorly  or  posteriorly.  The  body,  including  the  snout,  is 
otherwise  unmarked  black  (see  Stuart,  1943a,  plate  II,  fig. 
3).  The  eye  is  dark. 

Bolitoglossa platvdactyla  has  a single  broad  middorsal  light 
swath  on  a generally  dark  ground.  The  snout  is  dark  and 
speckled  with  light  flecks;  additional  flecking  may  occur  on 
the  fore  limbs.  A light  swath  begins  on  the  eyelids  and  extends 
almost  to  the  tip  of  the  tail  (see  Wake  and  Lynch,  1976,  fig. 
43;  Stuart,  1943a,  plate  II,  fig.  1);  the  eye  is  dark. 

Finally,  B.  salvinii  has  the  same  distribution  of  light  dorsal 
markings  as  B.  Jlaviventris'.  paired  dorsolateral  lines  that  join 
anteriorly  to  produce  a light-colored  snout  and  that  frequent- 
ly meet  across  the  dorsal  midline  or  break  into  reticulation. 
In  this  species,  the  ventral  surfaces  are  dark  and  undiffer- 
entiated from  the  dark  flanks,  although  a light  ventral  blotch 
occasionally  occurs  (see  Wake  and  Lynch,  1976,  fig.  24; 
Schmidt,  1936,  fig.  17).  The  eye  may  be  either  light  or  dark. 

Relationships  among  the  B.  mexicana  group  species  re- 
main obscure.  The  animals  are  scarce,  and  more  samples 
would  be  needed  for  a comprehensive  analysis  of  the  entire 
assemblage.  While  primitive-derived  polarities  are  not 
understood  for  the  color  characters  separating  the  species, 
the  seven  taxa  cluster  in  two  basic  color  and  pattern  groups. 
Stuart  (1943a)  considered  B.  platvdactyla,  B.  mexicana,  B. 
mulleri,  and  B.  odonelH  to  form  a group  united  by  aspects 
of  their  coloration,  and  the  remaining  three  taxa,  B.  jacksoni, 


B.  Jlaviventris,  and  B.  salvinii,  themselves  form  a cluster.  All 
three  of  the  latter  forms  have  ( 1 ) some  white  edging  between 
dark  and  light  areas;  (2)  at  least  some  individuals  with  golden 
eyes;  (3)  the  same  basic  hues,  yellow  (to  tan)  and  chocolate 
brown;  and  (4)  a dorsal  swath  of  brown  bordered  bilaterally 
and  on  the  snout  by  yellow.  These  three  species  are  the  only 
members  of  the  mexicana  group  that  ever  have  light  ventral 
coloration  (apparently  universal  in  B.  jacksoni  and  B.  jla- 
viventris and  occasionally,  as  small  blotches,  in  B.  salvinii). 
An  additional  unique  feature  of  the  jacksoni-jlaviventris-sal- 
vinii  assemblage  is  that  each  is  marked  only  with  solid  colors, 
the  light  regions  a single  uniform  yellow  and  the  dark  regions 
an  equally  pure  brown;  speckling  and  mottling  are  never 
observed. 

ETYMOLOGY.  I name  this  rare  and  beautiful  new  species 
for  Jeremy  L.  Jackson,  the  friend  who  aided  me  in  long  moldy 
months  of  wet  season  collecting,  and  the  captor  of  the  first, 
and  now  sole,  representative  of  this  species. 

Bolitoglossa  lincolni  (Stuart,  1943a) 

Figure  3 

Oedipus  lincolni  Stuart,  1943a:9. 

Magnadigita  lincolni:  Taylor,  1944:218. 

Bolitoglossa  lincolni:  Wake  and  Brame,  1963:386. 
Bolitoglossa  resplendens  McCoy  and  Walker,  1966:1. 

TYPE  LOCALITY.  Monte  at  Salquil  Grande,  Depto.  El 
Quiche,  Guatemala,  2,450  m elevation. 

RANGE.  Including  the  range  of  B.  resplendens  (herein 
synonymized  with  B.  lincolni ),  this  is  one  of  the  widest  rang- 
ing of  Central  American  salamanders.  It  occurs  on  the  Pacific 
Uplands,  in  the  Cuilco,  in  the  Cuchumatanes,  and  on  the 
Mesa  Central  de  Chiapas.  B.  lincolni  is  herein  reported  from 
one  new  locality  in  the  Cuchumatanes  and  one  in  the  Cuilco. 

HABITAT  AND  HABITS.  Bolitoglossa  lincolni  were  tak- 
en in  bromeliads  and  under  logs  and  bark  (see  Wake  and 
Lynch,  1976,  fig.  30).  In  addition  they  were  found  to  be  active 
on  wet  nights.  The  species  occurs  in  the  upper  subtropical 


Table  2.  Summary  of  geographic  variation  in  Bolitoglossa  lincolni. 


Population 

Dorsal 

pattern 

Ventral 

spotting 

Foot 

webbing 

Max.  standard 
length 

(millimeters)* 

Pacific  Uplands 

Swath  to  spots 

>95%  of  pop. 

Reduced 

m 

76 

f 

74 

Cuilco 

Swath  to  spots 

<60%  of  pop. 

Intermediate 

m 

58 

f 

79 

Cuchumatanes 

Swath 

>95%  of  pop. 

Extensive 

m 

66 

f 

78 

Mesa  Central 

Swath 

>95%  of  pop. 

Reduced 

m 

62.3** 

r 

78.4** 

* Males  (m),  females  (0. 

**  Data  from  McCoy  and  Walker  (1966). 


Contributions  in  Science,  Number  348 


Elias:  Guatemalan  Salamanders  9 


forest  from  well  above  the  cloud  line  up  to  the  lower  tem- 
perate zone. 

SYSTEMATICS.  The  diagnostic  characters  differentiat- 
ing Bolitoglossa  resp/endens  from  B.  lincolni  are  its  greater 
standard  length  and  proportionately  shorter  limbs.  In  the 
description  of  B.  resplendens  it  was  reported  that  a higher 
incidence  of  ventral  spotting  and  less  extensive  interdigital 
webbing  also  distinguish  B.  resplendens  from  B.  lincolni 
(McCoy  and  Walker,  1966). 

Examination  of  specimens  from  the  four  known  popula- 
tions indicates  that  the  two  supposedly  diagnostic  features 
of  B.  resplendens  reflect  comparison  to  a limited  sample  of 
unusually  small  B.  lincolni.  Nominate  B.  lincolni  from  the 
Cuchumatanes  are  fully  as  large  as  the  specimens  in  the  B. 
resplendens  type  series  (McCoy  and  Walker,  1966)  from  the 
Mesa  Central  de  Chiapas  (Table  2),  and  both  populations 
have  proportionately  longer  limbs  at  smaller  standard  length 
(Fig.  8).  Specimens  from  the  Cuilco  and  Pacific  Uplands  have 
the  same  maximum  size  (Table  2)  and  limb/body  length 
allometry  (Fig.  8)  as  the  other  two  populations. 

Color  pattern,  both  dorsal  and  ventral,  and  degree  of  in- 
terdigital webbing  vary  between  the  four  populations  (see 
Table  2 and  Fig.  3 for  dorsum  of  Cuchumatan  specimen)  but 
not  in  concordant  patterns  that  suggest  species  level  differ- 
entiation. Thus,  I refer  the  name  Bolitoglossa  resplendens  to 
the  synonymy  of  Bolitoglossa  lincolni. 

Wake  and  Lynch  (1982)  reported  the  results  of  morpho- 
metric and  electrophoretic  analysis  of  the  Bolitoglossa  frank- 
lini  species  group.  While  there  is  general  agreement  with  my 
conclusions,  they  choose  to  continue  to  recognize  Bolito- 
glossa resplendens  as  a distinct  species  based  upon  the  large 
genetic  distance  between  the  Cuilco  population  and  those 
from  the  Cuchumatanes  and  Pacific  Uplands.  The  genetic 
affinities  of  the  nominate  populations  of  Bolitoglossa  re- 
splendens on  the  Mesa  Central  de  Chiapas  remain  unknown. 

Bolitoglossa  morio  (Cope,  1869) 

Figure  3 

Geotriton  carbonarius  (part)  Cope,  1868:313. 

Oedipus  morio  Cope,  1869:103. 

Geotriton  morio:  Smith,  1877:64. 

Spelerpes  morio:  Boulenger,  1882:70. 

Spelerpes  bocourti  Brocchi,  1883:1  1 1. 

Magnadigita  morio:  Taylor,  1944:218. 

Magnadigita  omniumsanctorum  Stuart,  1952:4. 

Bolitoglossa  omniumsanctorum:  Wake  and  Brame,  1963:386. 
Bolitoglossa  morio:  Wake  and  Brame,  1963:386. 

TYPE  LOCALITY.  Mountains  of  Guatemala. 

RANGE.  This  species  occurs  throughout  the  Pacific  Up- 
lands, in  the  Chuacus,  and  in  the  Cuchumatanes.  It  is  here 
reported  for  the  first  time  from  the  Cuilco. 

HABITAT  AND  HABITS.  B.  morio  were  taken  under 
logs,  in  logs,  and  in  bromeliads  and  were  found  to  be  active 
on  rainy  nights.  They  occur  in  the  upper  subtropical  and 
lower  temperate  zone  forests  between  roughly  1,800  and 
3,000  m. 


A 


18- 

• 

• A 

OA  OO  «Of 

° 

• o 

• 2 

oO  A A o 

| 14- 

it  A • 

-C 

°o°» 

O'  _ 

c 

A* 

A A • 

Bolitoglossa  lincolni 

1 l0~ 

• 

o 

A Mesa  Central  de  Chiapas 

a Cuchumatanes 

c — 

A • 

X 

A 

• Cuilco 

6- 

• 

1 1 1 1 

O Pacific  uplands 

i ~T — r~ — ~n 

10  20  30  40  50 

Axilla -groin  length  (mm) 


Figure  8.  Relationship  between  hind  limb  length  and  axilla-groin 
length  in  four  populations  of  Bolitoglossa  lincolni. 

SYSTEMATICS.  Relationships  of  this  species  are  ob- 
scure, but  its  closest  relative  appears  to  be  B.  flavimembris 
(see  species  group  section,  below).  The  characters  that  unite 
the  populations  of  this  species  are  the  presence  of  large  light 
(whitish,  cream,  or  pink)  blotches  on  the  flanks  and/or  sides 
of  the  tail;  the  slate  grey  to  brownish  ground  color  of  the 
dorsum;  salt  and  pepper  belly  mottling;  reduced  webbing 
such  that  all  toes  of  the  hind  foot  are  free  at  their  tips;  and 
the  robust  body  form. 

The  newly  discovered  Cuilco  population  has  all  of  the 
specific  characters,  except  that  the  lateral  markings  are  re- 
duced to  small  flecks  (diameters  less  than  that  of  the  eye), 
which  are  restricted  to  the  sides  of  the  tail  rather  than  ex- 
tending onto  the  trunk. 

Bolitoglossa  mulleri  (Brocchi,  1883) 

Figure  5 

Spelerpes  mulleri  Brocchi,  1883:1 16. 

Oedipus  p/atydactylus  (part)  Dunn,  1926:400. 

Oedipus  mulleri:  Schmidt,  1936:150. 

Bolitoglossa  mulleri:  Taylor,  1944:219. 

Oedipus  mexicanus  mulleri:  Stuart,  1948:19. 

Bolitoglossa  moreleti  mulleri:  Duellman,  1963:220. 
Bolitoglossa  mexicana  mulleri:  Wake  and  Brame,  1963:386. 

TYPE  LOCALITY.  Several  localities  in  Alta  Verapaz,  re- 
stricted by  Stuart  (1943a)  to  "les  montagnes  qui  dominent 
Coban.” 

RANGE.  This  form  had  been  taken  only  in  the  Xucaneb 
complex  until  discovered  in  the  Cuchumatanes  during  the 
study  described  here.  It  is  now  known  from  Santa  Cruz  Bari- 
llas and  from  a finca  3 km  to  the  NE  at  1,500  m (Fig.  7). 

HABITAT  AND  HABITS.  This  species  occurs  in  the  low 
pine-broadleaf  cloud  forest  border  in  the  Xucaneb  and  was 
found  in  the  Cuchumatanes  in  the  cutover  Barillas  valley, 
which  was  once  a mix  of  the  two  forest  types.  This  area  is 
at  the  elevation  of  the  lower  subtropical  zone  but  is  drier 


10  Contributions  in  Science,  Number  348 


Elias:  Guatemalan  Salamanders 


because  of  its  protected  position.  Of  the  ten  or  so  specimens 
taken  from  the  Cuchumatanes,  all  but  one  were  found  under 
stones  in  a well-watered  garden  in  the  town  of  Barillas.  The 
other  specimen  was  found  under  a piece  of  wood  in  a stand 
of  coffee.  In  the  Xucaneb  the  species  has  been  taken  between 
1,000  and  1,500  m. 

SYSTEMATICS.  The  new  population  has  a somewhat 
reduced  middorsal  light  stripe  relative  to  animals  from  the 
type  locality:  markings  never  extend  anterior  to  the  nape, 
and  some  individuals  are  unmarked  black. 

Bolitoglossa  mulleri  is  a member  of  the  Bolitoglossa  mex- 
icana  species  group  (sensu  Wake  and  Lynch,  1 976).  Variation 
and  relationships  among  the  mexicana  group  species  are  dis- 
cussed in  the  B.  jacksoni  account  above. 

Bolitoglossa  rostrata  (Brocchi,  1883) 

Figure  3 

Spelerpes  rostratum  Brocchi,  1883:1  12. 

Oedipus  rostratum:  Dunn,  1924:99. 

Oedipus  rostratus:  Dunn,  1926:384. 

Magnadigita  rostrata:  Taylor,  1944:218. 

Bolitoglossa  rostrata:  Wake  and  Brame,  1963:386. 

TYPE  LOCALITY.  The  mountains  of  Totonicapan,  Gua- 
temala. 

RANGE.  This  species  is  one  of  the  most  broadly  ranging 
forms  in  Nuclear  Central  America,  from  the  Mesa  Central 
de  Chiapas  through  the  Cuchumatanes  and  into  the  Pacific 
Uplands.  A newly  discovered  population  at  the  eastern  ex- 
tremity of  the  Cuchumatanes  indicates  that  the  species  may 
be  distributed  throughout  that  cordillera. 

HABITAT  AND  HABITS.  B.  rostrata  were  taken  in  and 
under  logs,  under  bark,  in  bunchgrass,  and  under  rocks,  and 
were  found  to  be  active  on  rainy  nights.  The  species  occurs 
only  above  2,700  m and  up  to  about  3,000  m,  in  temperate 
forests  and  above  the  treeline  in  bunchgrass  associations. 

SYSTEMATICS.  The  species  is  distinguished  by  the  fol- 
lowing characters:  it  has  almost  unwebbed  toes  (as  unwebbed 
as  any  member  of  the  genus);  it  is  brown,  typically  with  a 
middorsal  swath  of  light  color  or  light  shoulder  stripes;  and 
it  has  a uniform  light  beige  to  dull  yellow  ventral  and  sub- 
caudal  color. 

B.  rostrata  appears  to  be  most  closely  related  to  the  B. 
dunni  series  (see  B.  cuchumatana  account  above)  and  where 
it  occurs  in  sympatry  with  B.  cuchumatana  the  two  species 
are  separated  with  difficulty.  Bolitoglossa  rostrata  is  distin- 
guishable from  B.  cuchumatana  on  a number  of  subtle  char- 
acters; it  occurs  in  an  unstriped  morph,  is  slightly  less  robust, 
has  a longer  tail,  and  shows  reduced  interdigital  webbing  (Fig. 

3). 

Bolitoglossa  rufescens  (Cope,  1869) 

Figure  5 

Oedipus  rufescens  Cope,  1869: 104. 

Geotriton  rufescens:  Smith,  1877:76. 

Spelerpes  rufescens:  Boulenger,  1882:71. 


Bolitoglossa  rufescens:  Taylor,  1 94 1 : 1 45. 

Palmatotriton  rufescens:  Smith,  1945:4. 

TYPE  LOCALITY.  Orizaba,  Veracruz,  Mexico. 

RANGE.  The  range  of  this  species  is  broad,  extending 
along  the  Atlantic  versant  throughout  western  Honduras, 
Guatemala,  and  Chiapas,  Mexico,  and  continuing  up  the 
Atlantic  coastal  plain  through  the  Mexican  state  of  Veracruz 
to  San  Luis  Potosi.  The  species  is  well  known  from  Chiapas 
and  the  low  Xucaneb  on  both  sides  of  the  Cuchumatanes, 
but  this  is  the  first  report  of  its  occurrence  in  the  Cuchu- 
matanes proper.  It  occurs  just  below  the  cloud  line  at  Finca 
Chiblac  in  the  northwestern  Cuchumatanes. 

HABITAT  AND  HABITS.  This  species  almost  always 
occurs  in  the  leaf  bases  of  banana  plants.  All  but  one  of  the 
four  B.  rufescens  taken  in  the  Cuchumatanes  were  found  in 
this  situation.  The  exception  was  found  crossing  a trail  on  a 
sultry,  overcast  afternoon  at  about  1 400  h.  The  species  occurs 
throughout  the  tropical  zone  forest  from  near  sea  level  (Stuart, 
1943a)  up  to  the  1,200-m  cloud  line. 

SYSTEMATICS.  Specimens  from  the  Cuchumatanes  were 
assigned  to  B.  rufescens  based  upon  derived  skull  and  foot 
structure  and  the  absence  of  maxillary  teeth. 

Bradytriton  silus  Wake  and  Elias,  1983 

Figure  9 

Bradytriton  silus  Wake  and  Elias,  1983:3. 

TYPE  LOCALITY.  Finca  Chiblac,  15  km  NE  Barillas, 
Huehuetenango,  Guatemala,  elevation  4,300  ft.  (1,310  m). 

RANGE.  This  species  was  discovered  in  the  course  of  this 
investigation.  It  is  known  only  from  the  type  locality. 

HABITAT  AND  HABITS.  Eleven  of  the  twelve  known 
specimens  were  taken  beneath  pieces  of  wood  in  a clearing 
that  was  being  prepared  for  the  cultivation  of  quinine.  The 
other  individual  was  found  under  a log  in  the  undisturbed 
subtropical  zone  cloud  forest  surrounding  the  clearing.  The 
native  forest  at  the  type  locality  is  a mixed  hardwood  growth 
that  receives  over  6 m of  rain  yearly. 

SYSTEMATICS.  This  species  has  unique  specializations 
and  has  been  placed  in  a monotypic  genus.  Its  possible  re- 
lationships are  discussed  by  Wake  and  Elias  (1983). 

Dendrotriton  cuchamatanus 
(Lynch  and  Wake,  1975) 

Figure  5 

Chiropterotriton  cuchumatanus  Lynch  and  Wake,  1975:6. 
Dendrotriton  cuchumatanus:  Wake  and  Elias,  1983. 

TYPE  LOCALITY.  On  highway  9N,  8.5  km  (by  road)  SW 
San  Juan  Ixcoy,  Depto.  Huehuetenango,  Guatemala,  eleva- 
tion 2,860  m. 

RANGE.  This  species  is  known  only  from  the  type  locality. 
Appropriate  habitat  is  present  elsewhere  in  the  area,  and  this 
species  may  range  more  widely  than  is  presently  known. 

HABITAT  AND  HABITS.  This  species  has  been  taken 
under  moss  and  bark  on  large  fallen  trees.  The  type  locality 
is  a patch  of  great  buttressed  rain  forest  oaks  on  the  Sub- 


Contributions  in  Science,  Number  348 


Elias:  Guatemalan  Salamanders  1 1 


Figure  9.  Bradytriton  silus,  Pseudoewycea  rex.  and  Nyctanolis  pernix,  life  size.  UPPER  LEFT:  Bradytriton  situs,  head,  legs,  and  feet  slate 
grey,  trunk  and  tail  brick  red-brown.  LOWER  LEFT:  Pseudoewycea  rex  (Cuchumatan  specimen),  dark  brown  overall  (in  some  Cuchumatan 
specimens  and  in  most  Cuileo  animals,  there  is  light  speckling,  especially  laterally).  RIGHT:  Nyctanolis  pernix.  background  color  black,  spots 
on  eyelids  and  nape  crimson,  those  on  elbows  and  knees  orange,  trunk  and  tail  spots  yellow. 


tropical-temperate  zone  border.  This  area  receives  a great 
but  undetermined  amount  of  precipitation  both  in  the  form 
of  rain  and  as  dense,  cold  fog. 


SYSTEMATICS.  The  genus  Dendrotriton  has  been  re- 
viewed by  Lynch  and  Wake  (1975).  Their  results  suggest  dose 
relationship  between  D.  cuchumatanus,  D.  bromeliacea,  and 


12  Contributions  in  Science,  Number  348 


Elias:  Guatemalan  Salamanders 


20- 


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if) 

c r> 
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o 


lo- 


ci 3 


A D cuchumatanus  (WC) 
3 D.  rabbi  (EC) 

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rorViAAYx  n A_ 


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50  60 

Braincase  width 


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70 


Figure  10.  Relationship  between  length  of  the  preorbital  process 
of  the  vomer  and  braincase  width  in  Dendrotriton  rabbi  and  D. 
cuchumatanus  (in  micrometer  units).  WC  = Western  Cuchuma- 
tanes;  EC  = Eastern  Cuchumatanes;  CU  = Cuilco. 


D.  megarhinus  (the  latter  two  are  inhabitants  of  the  Pacific 
Uplands),  based  upon  shared  configurations  of  the  vomer 
and  the  nostril. 

Dendrotriton  rabbi  (Lynch  and  Wake,  1975) 

Figure  5 

Chiropterotriton  rabbi  Lynch  and  Wake,  1975:2. 
Dendrotriton  rabbi:  Wake  and  Elias,  1983. 

TYPE  LOCALITY.  9.5  km  W,  8.5  km  S (by  air)  La  De- 
mocracia,  Depto.  Fluehuetenango,  Guatemala,  elevation 
2,100  to  2,500  m. 

RANGE.  This  species,  described  and  hitherto  known  only 
from  a spur  of  the  Montanas  de  Cuilco,  is  here  reported  both 
from  the  main  massif  of  the  Cuilco  and  from  two  localities 
in  the  Cordillera  de  los  Cuchumatanes. 

HABITAT  AND  HABITS.  Dendrotriton  rabbi  occur  at 
the  subtropical-temperate  border.  In  the  Cuilco,  they  have 
most  frequently  been  found  in  bromeliads  but  only  under 
bark  on  logs  and  stumps  in  the  Cuchumatanes,  perhaps  be- 
cause bromeliads  are  absent  from  the  appropriate  elevations. 
The  species  occurs  from  2,100  to  2,700  m. 

SYSTEMATICS.  The  newly  discovered  populations  have 
been  referred  to  D.  rabbi  on  the  basis  of  a small  nostril  di- 
ameter and  the  presence  of  a preorbital  process  on  the  vomer 
in  adults  (as  distinct  from  D.  cuchumatanus,  D.  brome/iacea, 
and  D.  megarhinus)  and  because  some  specimens  from  each 
population  have  ossified  septomaxillary  bones  (absent  in  D. 
xolocalcae)  (for  discussion,  see  Lynch  and  Wake,  1975,  1978). 

Lynch  and  Wake  (1975)  examined  the  ontogeny  of  the 
nostril  in  all  of  the  species  of  Dendrotriton  and  discovered 
that  all  juveniles  have  similar,  relatively  large  nostrils.  They 
found  that  patterns  of  relative  growth  determined  the  adult 


condition;  species  that  as  adults  have  small  nostrils  experi- 
ence either  no  growth  or  actual  shrinkage  in  the  absolute  size 
of  the  nostril  through  ontogeny,  while  large  nostriled  forms 
experience  nostril  growth. 

I have  examined  ontogenetic  variation  in  the  presence  or 
absence  of  the  preorbital  process  of  the  vomer.  The  process 
is  large  and  tooth-bearing  in  small  nostriled  species  but,  if 
present,  is  a nub  in  species  with  large  nostrils.  As  with  nostril 
size,  vomerine  process  length  is  initially  equal  in  the  large 
nostriled  D.  cuchumatanus  and  small  nostriled  D.  rabbi.  Dif- 
ferential growth  produces  the  difference  in  the  adult  condition 
in  the  two  species  (Fig.  10).  In  this  case,  interspecific  differ- 
ences are  compounded  by  a differential  in  maximum  adult 
size.  These  data  further  indicate  that  the  newly  discovered 
Cuchumatan  populations  clearly  fall  with  the  nominate  D. 
rabbi.  With  the  new  localities  reported  here,  D.  rabbi  is  known 
to  occur  within  250  vertical  m and  about  25  km  of  D.  cu- 
chumatanus in  the  Cuchumatanes.  The  two  species  have  been 
taken  in  similar  habitats  without  intervening  ecological  bar- 
riers, and  the  proximity  of  the  two  suggests  that  the  first  case 
of  sympatry  in  the  genus  may  yet  be  found. 

Nyctanolis  pernix  Elias  and  Wake,  1983 

Figure  9 

Nyctanolis  pernix  Elias  and  Wake,  1983:2. 

TYPE  LOCALITY.  Finca  Chiblac,  about  10  km  NE  Santa 
Cruz  Barillas,  Depto.  Huehuetenango,  1,370  m (4,500  ft.) 
elevation. 

RANGE.  This  species  has  been  taken  at  the  type  locality, 
just  across  the  Mexican  border  in  Chiapas  and  in  the  western 
end  of  the  Sierra  de  las  Minas  near  Purulha,  Baja  Verapaz 
(J.  Campbell,  pers.  comm.). 

HABITAT  AND  HABITS.  The  type  series  was  found 
under  moss  and  bark  during  the  day  and  on  stumps  and  logs 
on  wet  nights.  The  type  locality  is  a region  of  quinine  cul- 
tivation in  the  subtropical  zone,  just  above  the  cloud  line. 
The  forest  receives  over  6 m of  rain  annually.  The  Mexican 
specimen  was  taken  in  the  mouth  of  a cave  in  the  cloud 
forest. 

SYSTEMATICS.  This  species  was  discovered  during  the 
course  of  this  investigation  and  appears  to  have  no  close 
relatives.  The  relationships  of  this  monotypic  genus  were 
discussed  by  Elias  and  Wake  (1983). 

Pseudoeurycea  rex  (Dunn,  1921) 

Figure  9 

Oedipus  rex  Dunn,  1921:143. 

Pseudoeurycea  rex:  Taylor.  1944:209. 

TYPE  LOCALITY.  Sierra  Santa  Elena  (near  Tecpam), 
Guatemala,  elevation  9,500  ft.  (2,900  m). 

RANGE.  P.  rex  is  known  from  the  Cuchumatanes  and 
from  several  regions  on  the  Pacific  Uplands.  The  species  is 
here  recorded  for  the  first  time  from  the  Montanas  de  Cuilco. 

HABITAT  AND  HABITS.  This  form  occurs  from  2,750 
m elevation  to  over  4,000  m at  the  top  of  the  highest  moun- 


Contributions  in  Science,  Number  348 


Elias:  Guatemalan  Salamanders  13 


tains  in  its  range  and  occurs  in  the  temperate  zone  forest  and 
above  the  treeline  in  open  bunchgrass  communities.  They 
live  under  and  within  logs,  under  bark,  moss,  grass  clumps, 
and  stones  and  were  found  active  in  the  open  on  wet  nights. 
P.  rex  is  often  abundant,  reaching  densities  higher  than  those 
of  any  other  Guatemalan  salamander. 

SYSTEMATICS.  The  Guatemalan  members  of  the  genus 
Pseudoeurycea  are  four  or  five  similar  brown  to  black-colored 
species.  Species-level  differences  in  foot  webbing  and  color 
pattern  have  proven  difficult  to  document  and  tend  to  be 
exceeded  by  intrapopulational  variation.  No  careful  quan- 
titative work  on  variation  in  this  assemblage  has  been  pub- 
lished, and  my  allocation  of  the  Cuilco  and  Cuchumatan 
populations  is  based  upon  their  elevational  distribution  and 
their  general  morphological  similarities  to  known  popula- 
tions of  P.  rex. 

DISCUSSION 

SPECIES  GROUPS  IN  BOLITOGLOSSA  BETA 

Since  Wake  and  Lynch  (1976)  last  outlined  species  groups 
in  the  genus  Bo/itog/ossa,  examination  of  new  material  from 
the  Cuchumatanes,  the  Cuilco,  the  Pacific  Uplands,  and  the 
Sierra  de  las  Minas  has  blurred  certain  distinctions  and  clar- 
ified others  among  the  species  found  in  Nuclear  Central 
America  (abbreviated  NCA  and  including  the  entire  land- 
mass  between  the  Isthmus  of  Tehuantepec  and  the  Nicara- 
guan Depression).  Implicit  in  some  of  the  comparisons  made 
in  the  preceding  species  accounts,  and  crucial  to  the  discus- 
sion sections  to  come,  is  a revision  of  the  species  groups  for 
Guatemalan  Bolitoglossa. 

Wake  and  Dresner  (1967)  found  that  a group  of  derived 
structural  characters  involved  in  tail  autotomy  was  shared 
by  certain  species  of  Bolitoglossa.  The  derived  group,  referred 
to  by  Wake  and  Lynch  ( 1 976)  as  "Bolitoglossa  beta,”  includes 
all  Mexican  and  NCA  species  except  the  B.  mexicana  group, 
as  well  as  two  forms  from  the  Talamancan  highlands  further 
south.  Included  are  the  following  species:  B.  alvaradoi,  B. 
arboroscandens.  B.  cuchumatana,  B.  dofleini,  B.  dunni,  B. 
engelhardti,  B.  flavimembris,  B.  franklini,  B.  hartwegi,  B. 
helmrichi,  B.  lincolni,  B.  macrinii,  B.  melania,  B.  morio,  B. 
occidentalis,  B.  riletti.  B.  rostrata.  B.  rufescens,  B.  schmidti, 
B.  stuarti,  B.  veracrucis,  B.  yucatana.  and  at  least  one  un- 
described species  (Wake  and  Lynch,  1982). 

Species  groups  within  Bolitoglossa  beta  were  first  recog- 
nized informally.  Taylor  (1941),  in  his  description  of  B.  oc- 
cidentalis. noted  the  affinity  between  his  new  species  and  B. 
rufescens.  Stuart  (1943a)  noted  that  B.  dofleini  and  B.  yu- 
catana were  close  relatives.  He  also  suggested  that  B.  cuchu- 
matana, B.  dunni,  B.  engelhardti  and  B.  helmrichi  formed  a 
close  group  in  terms  of  both  morphology  and  ecology,  and 
in  a later  paper  (1952)  he  named  them  the  B.  dunni  species 
group.  In  the  description  of  B.  lincolni  Stuart  (1943a)  re- 
ported the  new  species  to  be  most  closely  affiliated  with  B. 
franklini  on  morphological  grounds,  and  later  (Stuart,  1952) 
included  them  with  the  then  newly  described  B.  nigroflaves- 
cens  in  a B.  franklini  species  group.  Recently  Wake  and  Lynch 
( 1 982)  reduced  B.  nigroflavescens  to  the  status  of  a subspecies 


within  B.  franklini.  Stuart  (1952)  erected  the  B.  morio  group 
to  contain  B.  morio  and  B.  omniumsanctorum,  but  Wake 
and  Elias  (1983)  synonymized  the  two. 

By  1952,  therefore,  five  species  assemblages  had  been  rec- 
ognized within  Bolitoglossa  beta,  leaving  unassigned  only  the 
following  six  of  the  nineteen  species  then  known:  B.  flavi- 
membris, B.  macrinii,  B.  riletti,  B.  rostrata,  B.  schmidti  and 
B.  veracrucis  (see  Table  3).  Wake  and  Brame  (1969)  named 
two  new  species,  B.  hartwegi  and  B.  stuarti,  and  demonstrat- 
ed their  close  morphological  similarities  to  one  another  and 
to  B.  veracrucis.  In  addition  they  reorganized  Stuart’s  ( 1 943a, 

1 952)  three  species  groups  into  two  new  ones,  the  B.  rostrata 
and  B.  helmrichi  groups  (Table  3).  These  two  groups  were 
large  and  heterogeneous  as  compared  to  those  described 
above.  The  B.  rostrata  group  included  Stuart’s  (1952)  B. 
morio  and  B.  franklini  groups,  B.  dunni  and  B.  engelhardti 
from  his  B.  dunni  group,  the  previously  unassigned  B.  ma- 
crinii, B.  riletti  and  B.  rostrata,  and  the  recently  described 
species  B.  resplendens  and  B.  brevipes.  Their  B.  helmrichi 
group  included  B.  cuchumatana  and  B.  helmrichi  from 
Stuart’s  (1952)  B.  dunni  group,  the  previously  unassigned  B. 
flavimembris  and  B.  veracrucis,  and  the  new  species  B.  hart- 
wegi and  B.  stuarti. 

Wake  and  Lynch  (1976)  assembled  the  first  complete  list- 
ing of  Bolitoglossa  species  groups.  They  tentatively  assigned 
every  species  to  some  group  and  recognized  a total  of  six 
species  groups  within  Bolitoglossa  beta.  The  B.  dofleini  group 
was  erected  to  include  B.  dofleini,  B.  yucatana  and  the  poorly 
known  B.  schmidti.  The  B.  rufescens  group  was  recognized 
as  containing  B.  occidentalis  and  B.  rufescens.  The  two  Tala- 
mancan species  from  southern  Central  America,  B.  alvaradoi 
and  B.  arboroscandens,  were  placed  together  in  a B.  alvaradoi 
group,  and  the  two  species  from  north  of  the  Isthmus  of 
Tehuantepec,  B.  macrinii  and  B.  riletti,  were  put  in  a B. 
macrinii  group.  The  two  large  groups,  B.  rostrata  and  B. 
helmrichi,  of  Wake  and  Brame  (1969),  were  retained  with 
their  original  membership  but  for  the  switching  of  B.  mono 
from  one  to  the  other  and  the  inadvertent  omission  of  B. 
omniumsanctorum  altogether  (see  Table  3). 

The  recognition  of  polyphyletic  groups  (groups  united  by 
characters  not  shared  by  their  most  recent  common  ancestor) 
is  a severe  obstacle  to  phylogenetic  reconstruction.  Ensuring 
that  only  nonpolyphyletic  (=  paraphyletic  and  monophyletic) 
species  groups  are  recognized  is  a first  step  towards  an  un- 
derstanding of  the  phylogenetic  history  of  Bolitoglossa  beta. 
To  eliminate  possibly  polyphyletic  groups  I have  recognized 
only  tight  clusters  of  phenotypically  similar  species.  Mor- 
phological similarity  between  species  group  members  is  dem- 
onstration of  nonpolyphyletic  origin  of  the  group  insofar  as 
such  similarities  are  homologous  (acquired  by  descent  from 
a similarly  endowed  common  ancestor). 

The  B.  rostrata  and  B.  helmrichi  groups  of  Wake  and  Lynch 
(1976)  are  large,  heterogeneous  and  united  by  no  specific 
characters.  To  compose  tighter  units  more  comparable  to 
other  species  groups  in  Bolitoglossa  beta  I have  subdivided 
and  reorganized  these  two  groups. 

The  B.  rostrata  group  contains  a series  of  species  that  has 
been  recognized  as  a phenetic  cluster  by  many  authors  (Stuart, 


14  Contributions  in  Science,  Number  348 


Elias:  Guatemalan  Salamanders 


Table  3.  The  species  groups  of  Bolitoglossa  beta  as  defined  by  different  authors.  Species  not  assigned  to  a group  until  after  1952  (both  newly  described  and  described  but  previously 
unaffiliated  species)  are  indicated  by  an  asterisk  where  they  first  appear. 

Early  classifications  Wake  and  Brame,  1969  Wake  and  Lynch,  1976  Elias,  1983 


Contributions  in  Science,  Number  348 


Elias:  Guatemalan  Salamanders  15 


group  I arbor oscande  ns 


1943a,  1952,  as  his  B.  frank/ini  group;  McCoy  and  Walker, 
1966,  Wake  and  Brame,  1969,  and  Wake  and  Lynch,  1976, 
as  the  B.  lincolni  subgroup  of  their  B.  rostrata  group;  Wake, 
Yang,  and  Papenfuss,  1980,  and  Wake  and  Lynch,  1982,  as 
the  B.  franklini  species  group).  This  assemblage  includes  B. 
franklini,  B.  lincolni.  B.  melania,  and  one  undescribed  Chia- 
pan  form  (Wake  and  Lynch,  1982,  who  also  include  B.  re- 
sp/endens).  These  species  are  characterized  by  large  adult  size, 
stout  bodies  and  tails,  and  a black  ground  color  broken  dor- 
sally  only  by  a sharply  bounded  swath,  spotting,  or  reticu- 
lation in  some  lighter  color;  red  in  some  species  to  yellow  or 
grey-green  in  others  (see  B.  lincolni.  Fig.  3).  B.  melania  is 
solid  black.  These  species  all  have  toetips  free  of  webbing 
and  subdigital  pads.  They  are  distributed  allopatrically  ex- 
cept for  an  area  of  elevational  parapatry  with  occasional 
hybridization  between  the  high  elevation  B.  lincolni  and  low- 
er elevation  B.  franklini  on  the  Pacific  Uplands  (Wake,  Yang, 
and  Papenfuss,  1980). 

The  remainder  of  the  B.  rostrata  group  of  Wake  and  Lynch 
(1976),  including  B.  dunni,  B.  engelhardti  and  B.  rostrata. 
fits  with  B.  cuchumatana  and  B.  helmrichi  of  their  B.  helm- 
richi group  to  form  an  assemblage  characterized  by  blunt 
rounded  toetips,  fully  developed  subdigital  pads,  and  a dark 
brown  ground  color,  frequently  marked  with  a lighter  brown 
dorsal  swath  or  paired  shoulder  stripes  (see  B.  rostrata  in 
Fig.  3 for  typical  color  pattern  and  compare  B.  rostrata  and 
B.  cuchumatana  [inset]  feet  as  examples  of  variation  in  foot 
morphology  within  the  group).  Stuart  (1943a,  1952)  recog- 
nized this  assemblage  as  his  B.  dunni  group  but  hesitated  to 
include  B.  rostrata  because  of  its  different  ecology.  Relying 
on  morphological  similarities,  I recognize  this  assemblage, 
including  B.  rostrata,  as  a new  B.  dunni  species  group.  The 
members  of  the  B.  dunni  group  are  all  either  geographically 
or  elevationally  (in  the  case  of  B.  rostrata  and  B.  engelhardti 
in  the  Pacific  Uplands)  allopatric,  except  that  narrow  sym- 
patry  is  seen  between  the  high  elevation  B.  rostrata  and  the 
lower  B.  cuchumatana  in  the  Cuchumatanes. 

Bolitoglossa  hartwegi.  B.  stuarti,  and  B.  veracrucis  were 
recognized  by  Wake  and  Brame  (1969)  to  form  a compact 
subgroup  within  their  B.  helmrichi  group.  I choose  to  rec- 
ognize the  similarities  of  these  species  by  isolating  them  in 
a new  B.  veracrucis  species  group.  The  three  species  are  united 
by  the  combination  of  fully  webbed  feet  that  lack  subdigital 
pads  and  are  differentiated  from  members  of  the  B.  rufescens 
group  by  posession  of  a strongly  articulated  skull  with  a broad 
pars  dentalis  of  the  premaxilla.  The  B.  veracrucis  group  has 
the  broadest  elevational  range  of  any  group  of  NCA  sala- 
manders (see  elevational  distribution  section  below),  and  no 
two  species  occur  sympatrically. 

The  two  remaining  species  left  over  from  Wake  and  Lynch’s 
(1976)  B.  helmrichi  group  are  B.  flavimembris  and  B.  morio. 
B.  morio  is  slightly  larger  than  B.  flavimembris,  but  the  two 
share  a stout  body  and  tail  and  a unicolor  dorsum  (see  B. 
morio  in  Fig.  3).  That  B.  flavimembris  is  not  affiliated  with 
the  B.  dunni  group  is  indicated  both  by  its  coloration  and 
build  and  by  its  rather  pointed  toetips  and  poorly  developed 
subdigital  pads  (see  Wake  and  Lynch,  1976,  fig.  22).  New 
evidence  that  B.  morio  and  B.  flavimembris  are  relatives 


comes  from  the  discovery  of  populations  of  intermediate 
morphology  in  the  Pacific  Uplands  of  Guatemala  near  the 
Mexican  border  (D.B.  Wake,  pers.  comm.).  Given  these  in- 
dications of  relationship,  and  the  absence  of  other  affilia- 
tions, I group  B.  flavimembris  and  B.  morio  together  in  a 
new  B.  morio  species  group. 

Aside  from  the  changes  proposed  above  I follow  the  species 
group  assignments  of  Wake  and  Lynch  (1976);  the  full  history 
of  species  groups  among  the  species  of  Bolitoglossa  beta 
including  my  revisions  is  presented  in  tabular  form  in 
Table  3. 

PATTERNS  OF  DISTRIBUTION  AND 
DIFFERENTIATION 

Faunal  Connections  Between  the  Mexican  Plateau, 

NCA,  and  the  Talamancas 

Wake  and  Lynch  (1976)  found  that  the  three  most  moun- 
tainous areas  of  Middle  America  are  especially  high  in  sal- 
amander species  diversity;  these  are  the  Caribbean  escarp- 
ment of  the  Mexican  Plateau,  the  mountains  of  NCA,  and 
the  Talamancan  highlands  of  Costa  Rica  and  Panama.  That 
there  has  been  faunal  contact  between  these  centers  of  di- 
versity is  demonstrated  by  the  distribution  of  some  genera 
on  more  than  one  of  the  three  upland  areas.  For  example, 
the  NCA  fauna  includes  some  genera  that  range  to  the  Mex- 
ican Plateau  ( Pseudoeurycea ),  others  that  range  to  the  Tala- 
mancas ( Oedipina  and  Nototriton ),  still  others  that  occur  on 
all  three  upland  areas  (Bolitoglossa),  as  well  as  genera  en- 
demic to  NCA  ( Bradytriton , Dendrotriton,  and  Nyctanolis). 
Each  genus  has  a clear  center  of  intrageneric  diversity  on  one 
of  the  three  upland  regions  (except  Bolitoglossa,  in  which  the 
alpha  and  beta  subgeneric  groups  center  on  two  different 
uplands  (Wake  and  Lynch,  1976)).  This  pattern  suggests  that 
most  intrageneric  evolution  proceeded  on  single  upland  areas 
at  a time  when  the  three  major  uplands  were  isolated  from 
one  another. 

If  we  go  one  step  further  back,  however,  and  consider  the 
origins  of  the  genera  themselves,  we  find  that  genera  with 
distributions  centered  on  the  same  upland  area  (i.e.,  Bradytri- 
ton, Dendrotriton,  Nyctanolis,  and  the  beta  complex  of  Bo- 
litoglossa in  NCA)  do  not  usually  form  a clearcut  monophy- 
letic  group  (Wake  and  Elias,  1983).  This  implies  a complex 
history  of  ancient  faunal  contact  between  the  three  great  up- 
land regions  of  Middle  America. 

Faunal  Connections  Among  the  Mountain  Ranges  of  NCA 

Within  NCA,  the  uplands  form  discrete  islands  of  habitat 
for  high  elevation  salamander  species.  Between  these  uplands 
lie  warmer,  drier,  rain-shadowed  valleys  that  are  generally 
uninhabited  by  salamanders.  Around  the  coastward  periph- 
ery of  these  mountains  are  warm,  humid  lowlands  that  sup- 
port a restricted  plethodontid  fauna.  Most  of  the  salamanders 
occurring  in  NCA  are  montane  (including  Bradytriton,  Den- 
drotriton, Nototriton,  Nyctanolis,  and  Pseudoeurycea).  A sin- 
gle genus,  Oedipina,  is  restricted  to  the  lowlands.  Only  Bolito- 
glossa occurs  at  all  elevations,  and  even  this  genus  is  split 


16  Contributions  in  Science,  Number  348 


Elias:  Guatemalan  Salamanders 


1000 


CUILCO 


CUCHUMATANES 


Figure  1 1 . Elevational  distribution  of  salamander  species  on  the  Montanas  de  Cuilco  and  the  Cordillera  de  los  Cuchumatanes  (all  collections 
combined). 


into  upland  ( B . helmrichi,  B.  morio,  B.  franklini)  and  low- 
land ( B . mexicana,  B.  dofleini,  B.  rufescens)  species  assem- 
blages (the  B.  veracrucis  group  is  exceptional  in  its  broad 
elevational  range  and  will  be  discussed  separately  below). 
The  upland  and  lowland  faunas  intermingle  narrowly  in  the 
low  cloud  forest  at  about  1 ,000  m elevation,  but  the  upland 
groups  are  subject  to  special  biogeographic  limitations,  which 
give  their  current  patterns  of  differentiation  particular  his- 
torical interest. 

Comparison  of  patterns  of  upland  diversity  obtaining  in 
the  Cuchumatanes  relative  to  the  Pacific  Uplands  raises  in- 
teresting contrasts.  There  are  fifteen  upland  salamander  species 
known  from  the  Pacific  Uplands  (Wake  and  Lynch,  1976, 
1982)  and  only  nine  (excluding  B.  hartwegi  of  the  B.  vera- 
crucis group)  from  the  Cuchumatanes,  but  the  supraspecific 
diversity  is  greater  in  the  Cuchumatanes.  The  nine  Cuchu- 
matan  species  belong  to  five  genera,  and  the  four  Bolitoglossa 
fall  into  three  species  groups.  In  the  Pacific  Uplands,  how- 
ever, the  fifteen  species  are  in  three  genera,  and  all  seven 
Bolitoglossa  are  in  the  same  three  species  groups  that  occur 
in  the  Cuchumatanes.  No  genus  or  species  group  in  the  Pacific 
Uplands  is  not  also  represented  in  the  Cuchumatanes,  but 
two  genera  present  in  the  Cuchumatanes  are  absent  in  the 
Pacific  Uplands.  The  two  areas  have  four  species  in  common: 


Bolitoglossa  lincolni  (here  including  B.  resplendens),  B.  mo- 
rio, B.  rostrata,  and  Pseudoeurycea  rex. 

The  fact  that  salamander  diversity  is  so  deep  in  the  Cu- 
chumatanes may  reflect  that  region's  ancient  history  as  a 
land-positive  area  (Rosen,  1978). 

The  shallowness  of  salamander  diversity  on  the  Pacific 
Uplands  may  reflect  the  topography  of  that  area:  the  high- 
lands are  composed  of  many  abutting  cinder  cones.  Small 
climatic  fluctuations  should  have  caused  alternate  fragmen- 
tation and  reunion  of  the  geographic  ranges  of  highland  sal- 
amander species.  When  a species’  range  is  subdivided  and 
then  reunified  after  a period  in  this  way,  a complex  of  in- 
trogressions,  parapatries  and  sympatries  might  be  expected 
to  result  as  partially  differentiated  sister  populations  are 
thrown  back  into  contact  after  a period  of  allopatry  on  neigh- 
boring cinder  cones.  Just  these  sorts  of  interactions  are  found 
among  the  Pacific  Uplands  salamanders  (Wake  and  Lynch, 
1976;  Wake,  Yang,  and  Papenfuss,  1980;  Wake  and  Lynch, 
1982). 

ELEVATIONAL  DISTRIBUTION 

Schmidt  (1936)  documented  the  elevational  ranges  of  sala- 
mander species  along  a transect  down  the  coastal  escarpment 


Contributions  in  Science,  Number  348 


Elias:  Guatemalan  Salamanders  17 


of  the  Pacific  Uplands.  He  found  that  species  there  inhabited 
narrow,  sharply  bounded  elevational  ranges  and  that  a given 
species  typically  inhabited  similar  elevations  at  different 
transect  sites  along  the  Pacific  escarpment. 

Wake  and  Lynch  (1976)  performed  a detailed  analysis  of 
elevational  distribution  along  a transect  near  the  volcano  first 
studied  by  Schmidt.  They  corroborated  Schmidt’s  results  and 
filled  them  out  with  detailed  studies  of  elevational  boundary 
zones  and  possible  interspecific  competition. 

Composite  transects  (grouping  all  collections)  for  the  Cu- 
chumatanes  and  Cuilco  (Fig.  1 1)  compared  to  the  transect 
results  of  Wake  and  Lynch  (1976)  for  Volcan  Tajumulco  in 
the  Pacific  Uplands  indicate  that  related  populations  (either 
conspecifics  or  members  of  related  species)  show  similar  el- 
evational ranges  wherever  they  occur.  Many  of  the  related 
populations  that  have  significant  variation  in  elevational  dis- 
tribution between  transects  appear  to  vary  in  response  to 
geographic  variation  in  the  elevation  of  the  cloud  line. 

The  results  for  the  Cuchumatanes  and  Cuilco  reinforce  the 
generalization  that  salamander  populations  in  NCA  inhabit 
narrow  elevational  belts,  and  recent  information  for  all  up- 
lands in  NCA  indicates  that  with  one  exception  no  single 
species  or  group  of  related  species  (meaning  species  groups 
in  Bolitoglossa  and  entire  genera  in  other  cases)  occurs  over 
an  elevational  range  as  broad  as  1,300  m.  The  exception  is 
Bolitoglossa  hartwegi  (with  a 1 ,500-m  elevational  range)  and 
the  Bolitoglossa  veracrucis  species  group  with  a combined 
elevational  range  of  over  2,800  m.  As  Wake  and  Brame 
(1969)  have  pointed  out,  and  as  these  results  reiterate,  this 
exceptional  group  may  have  played  a key  role  in  the  invasion 
of  the  lowlands  by  Bolitoglossa  beta  and  must  have  unique 
physiological  attributes,  which,  if  studied,  might  expand  our 
understanding  of  the  history  of  salamanders  throughout  Mid- 
dle America. 


SPECIMENS  EXAMINED 

Salamanders  are  known  from  the  following  localities  in  the 
Cordillera  de  Los  Cuchumatanes,  Depto.  Huehuetenango, 
and  the  Sierra  de  Cuilco,  Depto.  Quiche,  Guatemala. 

Bolitoglossa  cuchumatana.  CUCHUMATANES.  Finca 
Chiblac.ca.  10  km  (airline)  NE  Santa  Cruz  Barillas,  el.  1,300- 

l, 500  m,  3 km  S of  buildings,  MVZ  134524-72,  134574- 
79,  134582-88,  LACM  135482-85;  El  Porvenir,  134603- 
lb;  Finca  Chiblac,  149306-14;  1.5  km  S of  buildings,  134591- 
602;  San  Isidro;  134617-22;  El  Rayo,  134623-25.  On  ridge 
ca.  4 km  (airline)  NW  Santa  Cruz  Barillas,  el.  2,000  m,  MVZ 
149315. 

CUILCO.  6.7  mi.  (rd.)  S Nebaj  on  Rta.  Depto.  3,  el.  2,500 

m,  MVZ  160993-1005.  On  ridge  2.5  km  W,  2 km  N (airline), 
San  Miguel  Uspantan,  el.  2,200-2,500  m,  MVZ  149301-05. 
Oak  forest  about  2 km  (airline)  N Nebaj,  el.  1,900  m,  UMMZ 
89110  (holotype),  8911 1-13. 

Bolitoglossa  hartwegi.  CUCHUMATANES.  Finca  Chi- 
blac,  ca.  10  km  (airline)  NE  Santa  Cruz  Barillas,  el.  1,300- 


1.500  m,  MVZ  134629,  134631-33,  LACM  135481;  3 km 
S of  buildings,  MVZ  1 49324.  3 km  (airline)  NNW  San  Mateo 
Ixtatan,  el.  2,750  m,  MVZ  160355-59. 

CUILCO.  On  ridge  2.5  km  W,  2 km  (airline)  N San  Miguel 
Uspantan,  el.  2,200-2,500  m,  MVZ  149325. 

Bolitoglossa  jacksoni.  CUCHUMATANES.  Finca  Chi- 
blac,  ca.  12  km  (airline)  NE  Santa  Cruz  Barillas,  el.  1,400 
m,  MVZ  134634  (holotype). 

Bolitoglossa  lincolni.  CUCHUMATANES.  Just  N San  Juan 
Ixcoy,  UMMZ  123275-76.  Ca.  9 mi.  (rd.)  SW  El  Reposo, 
summit  between  El  Reposo  and  Oja  Blanca,  el.  2,200-2,300 
m,  MVZ  161791,  119011.  Planes  de  Pena  Blanca,  1.5  km 
(airline)  NE  Pena  Blanca  peak,  el.  2,700  m,  MVZ  149359- 
64;  9.5  km  W,  8.5  km  (airline)  S,  La  Democracia,  el.  2,100- 

2.500  m,  MVZ  103834-38;  10  km  W,  143681-87.  Paraiso 
Hoja  Blanca  Cumbre,  UMMZ  129149-50. 

CUILCO.  Monte  at  Salquil  Grande,  el.  2,450  m,  UMMZ 
89107  (holotype),  89108-09.  On  ridge  NW  San  Miguel  Us- 
pantan, el.  2,200-2,500  m,  2.5  km  N Uspantan,  MVZ 
149365-69;  3.5  km  (air)  NNW,  160833-39. 

Bolitoglossa  morio.  CUCHUMATANES.  Planes  de  Pena 
Blanca,  1.5  km  (airline)  NE  Pena  Blanca  peak,  el.  2,700  m, 
MVZ  149335-43.  Todos  Santos,  el.  2,500  m,  UMMZ 
102285-86  (holotype  and  paratype  of  Oedipus  omnium- 
sanctorum). 

Bolitoglossa  mulleri.  CUCHUMATANES.  Town  of  Santa 
Cruz  Barillas,  el.  1,500  m,  MVZ  131691-93,  149327-31, 
161 102.  Finca  la  Florida  ca.  2 mi.  (rd.)  NE  Santa  Cruz  Ba- 
rillas, el.  1,500  m,  MVZ  149326. 

Bolitoglossa  rostrata.  CUCHUMATANES.  Ca.  37-40  km 
(rd.)  N Huehuetenango,  el.  2,860  m,  MVZ  11341 8-540.  Vi- 
cinity Capzin,  Rta.  Nac.  9N,  50-60  km  (rd.)  N Huehuete- 
nango, el.  2,800-3,200  m.  52.7  km  N,  MVZ  114722-  65;  ca. 
50  km  N,  km  post  139,  115494-524,  117039.  2-8  km  N, 
W,  and  S (airline)  San  Mateo  Ixtatan,  el.  2,700-3,200  m;  2 
km  WSW,  MVZ  150940-160299;  6 km  WSW,  160300-333; 
5 km  SSW,  160334-49;  7 km  SSW,  160350-51;  3 km  NNW. 
160353,  160360;  4 km  NW,  163928-31,  163973;  2 km  W, 
171198-202;  4 km  SW,  171220-41;  4 km  N,  171242;  4 km 
W,  171173-97;  4 km  NW,  171112-43;  5 km  SW,  171215; 
5 km  S,  171203-14;  6 km  W,  171163-72;  8 km  W,  171144- 
62.  Paquix-Todos  Santos  road,  4-7  km  SE  Todos  Santos, 
MVZ  149198-206,  108616-30,  108631-72.  13.6  mi.  (rd.)N 
Santa  Eulalia  along  Rta.  Nac.  9N,  MVZ  1 49228.  5. 1 mi.  (rd.) 
N Santa  Eulalia  along  Rta.  Nac.  9N,  MVZ  108673-95, 
109296.  Cumbre  between  Ixtiapoc  and  Villa  Linda,  el.  2,800- 
3,400  m,  UMMZ  120007-08.  Cumbre  between  Yaiquich 
and  San  Mateo  Ixtatan,  el.  3,000  m,  UMMZ  120009.  Todos 
Santos,  el.  9,000-10,000  ft.,  UMMZ  120465-66,  120487. 
Mina  Villa  Linda,  UMMZ  130073.  9 km  (rd.)  SW  San  Juan 
Ixcoy,  el.  2,900  m,  UMMZ  120011.  Rta.  Nac.  9N,  7.4  mi. 
(rd.)  SW  San  Juan  Ixcoy,  MVZ  108696-728.  Rta.  Nac.  9N, 
10.6  mi.  (rd.)  SW  San  Juan  Ixcoy,  MVZ  108729-32.  8 km 
W San  Mateo  Ixtatan,  el.  3,100  m,  MVZ  171  148-62.  3 km 
NNW  San  Mateo  Ixtatan,  el.  8,700  ft.,  MVZ  150940-1000, 
160201-91,  160353,  160360;  4 km  NW,  2,750  m,  171112- 
43.  2 km  WSW  San  Mateo  Ixtatan,  el.  8,850  ft.,  MVZ  160292- 


18  Contributions  in  Science,  Number  348 


Elias:  Guatemalan  Salamanders 


99;  2 km  W,  2,750  m,  1 7 1 198-202.  6 km  WSW  San  Mateo 
Ixtatan,  el.  9,150-9,450  ft.,  MVZ  160330-33,  160352, 
160354;  6 km  W,  171  163-72.  5 km  SSW  San  Mateo  Ixtatan, 
el.  8,650-9, 1 50  ft.,  MVZ  1 60343-49;  4 km  SW,  2,835-2,965 
m,  171220-41;  4 km  W,  2,900  m,  171  173-97;  5 km  SW, 
2,965-3,045  m,  171215-19;  5 km  S,  3,150  m,  171203-14. 
7 km  SSW  San  Mateo  Ixtatan,  el.  9,650  ft.,  MVZ  160350- 
51.4  km  NW  San  Mateo  Ixtatan,  el.  2,750  m,  MVZ  1 63928— 
31,  163973;  4 km  N,  2,650  m,  171242. 

CUILCO.  Above  Salquil  Grande,  el.  3,000  m,  UMMZ 
891 16-18.  6.7  mi.  (rd.)  S Nebaj  by  Rta.  Depto.  3,  el.  2,500 
m,  MVZ  160992.  On  ridge  2.5  km  W,  2 km  (airline)  N San 
Miguel  Uspantan,  el.  2,260-2,640  m,  MVZ  149229-300. 

Bolitoglossa  rufescens.  CUCHUMATANES.  Vicinity  Fin- 
ca  Chiblac,  ca.  10  km  (airline)  NE  Santa  Cruz  Barillas,  el. 
950-1,100  m,  MVZ  134626-28. 

Bolitoglossa  rufescens.  CUCHUMATANES.  Vicinity  Fin- 
ca  Chiblac,  ca.  10  km  (airline)  NE  Santa  Cruz  Barillas,  el. 
950-1,100  m,  MVZ  134626-28. 

Bolitoglossa  stuarti  (tentative  identification).  CUCHU- 
MATANES. 15  km  (airline)  W La  Democracia,  el.  1,850  m, 
UMMZ  126781. 

Bradytriton  silus.  CUCHUMATANES.  Finca  Chiblac,  15 
km  NE  Barillas,  el.  4,300  ft.  (1,3 10  m),  MVZ  131586,  131587 
(holotype),  131589-94,  134635-37,  134638,  1 73063-64; 
LACM  134566. 

Dendrotriton  cuchumatanus.  CUCHUMATANES.  Along 
Rta.  Nac.  9N,  8.5  km  (rd.)  SW  San  Juan  Ixcoy,  el.  2,860  m, 
MVZ  113002  (holotype),  1 13003-05,  113007-19,  113021- 
22;  LACM  105296-97. 

Dendrotriton  rabbi.  CUCHUMATANES.  Planes  de  Pena 
Blanca,  1.5  km  (airline)  NE  Pena  Blanca  peak,  el.  2,700  m, 
MVZ  105298-300,  149702-05,  149139.  2 km  (airline)  WSW 
San  Mateo  Ixtatan,  el.  8,850  ft.,  MVZ  160437.  3 km  (airline) 
NNW  San  Mateo  Ixtatan,  el.  2,750  m,  160422-36.  Cloud 
forest  above  summit  of  road  between  El  Reposo  and  Oja 
Blanca  ca.  9 mi.  by  rd.,  SW  El  Reposo,  el.  2,200-2,300  m, 
MVZ  109297-301. 

CUILCO.  9.5  km  W,  8.5  km  S (airline),  La  Democracia, 
Montanas  de  Cuilco,  el.  2,100-2,500  m,  MVZ  1 14766-96, 
143920-26,  103839  (holotype),  103840-78,  LACM  105298- 
300.  On  ridge  2.5  km  W,  2 km  (airline)  N,  San  Miguel  Us- 
pantan, el.  2,400  m,  MVZ  160895-96. 

Nyctanolis pernix.  CUCHUMATANES.  Finca  Chiblac,  10 
km  (air)  NE  Barillas,  el.  1,370  m (4,500  ft.),  MVZ  131583- 
85,  134639-40,  134641  (holotype),  134642-44,  149370-73; 
MCZ  100154. 

Pseudoeurycea  rex.  CUCHUMATANES.  Vicinity  El  Re- 
tiro and  Pena  Blanca  peak,  el.  2,800-3,300  m,  MVZ  149404- 
33,  149464-520,  149522-58,  149693-700.  Vicinity  Capzin, 
Rta.  Nac.  9N,  50-60  km  (rd.)  N Huehuetenango,  el.  2,900- 
3,300  m,  MVZ  160973-82,  134211-401.2-8  km  N,  W,  and 
S (airline)  San  Mateo  Ixtatan,  el.  2,750-3,200  m;  5 km,  MVZ 
171243-52;  7 km  SSW,  160410-19;  6 km  WSW,  160409;  4 
km  NW,  167796-803.  Between  Cumbre  Yaiquich  and  San 
Mateo  Ixtatan,  3,000  m,  UMMZ  120067.  10  km  (rd.)  SW 
San  Juan  Ixcoy,  el.  3,300  m,  UMMZ  120068. 


CUILCO.  Above  Salquil  Grande,  el.  3,000  m,  UMMZ 
89114-15. 

ACKNOWLEDGMENTS 

I dedicate  this  work  to  the  memory  of  Lie.  Mario  Dary,  whose 
assistance  during  the  field  work  was  invaluable,  and  whose 
killing  was  tragic  and  senseless. 

Dr.  L.C.  Stuart  was  generous  with  his  hospitality  and  un- 
surpassed knowledge  of  Guatemala’s  herpetofauna.  The  res- 
idents of  the  departments  of  Huehuetenango  and  El  Quiche 
were  universally  tolerant  of  behavior  that  to  them  appeared 
absurd  and  were  friendly  and  hospitable  at  all  times.  The 
staffs  of  various  fincas,  especially  R.  Klein-Holkenborg  and 
Antonio  Diaz  of  Finca  Chiblac,  were  very  cooperative. 

A variety  of  other  collectors  assisted  me,  either  directly  in 
the  field  or  indirectly  through  the  sharing  of  experience  and 
specimens.  Among  these  are  J.L.  Jackson,  J.  Johnson,  E.J. 
Koford,  J.F.  Lynch,  T.J.  Papenfuss.  R.  Seib,  H.B.  Shaffer, 
L.C.  Stuart,  R.T.  Tesucun,  and  D.B.  Wake. 

The  students  and  staff  of  the  Museum  of  Vertebrate  Zo- 
ology provided  an  atmosphere  for  discussion  and  research, 
from  which  I profited  immensely.  Dr.  David  B.  Wake  ex- 
pedited every  aspect  of  this  research.  He  guided  the  inves- 
tigation through  its  entire  development,  provided  space  and 
resources  in  Berkeley,  and  was  the  source  of  most  field  sup- 
port (current  grant  NSF  DEB  78-03008). 

I should  finally  like  to  thank  Gene  M.  Christman  and 
James  Hendel  of  the  University  of  California  for  their  expert 
figure  preparation  and  K.F.  Liem  and  F.A.  Jenkins,  Jr.,  of 
Harvard  University,  under  whose  auspices  I received  support 
and  travel  funds  (on  NIH  Musculo-Skeletal  Training  Grant 
#5  T32  GMO  71 17-04  and  -05)  during  the  preparation  of 
this  manuscript. 

The  Spanish  summary  was  prepared  by  Fabian  Jaksic. 


LITERATURE  CITED 

Boulenger,  G.A.  1882.  Cat.  Batrach.  Grad.  British  Mu- 
seum, London.  127  pp.,  9 plates. 

Brocchi,  P.  1 883.  Etude  des  batraciens  de  I’Amerique  Cen- 
trale.  Mission  Scientifique  au  Mexique  et  dans  l’Ame- 
rique  Centrale  3(2):  1-1 22,  21  plates. 

Cope,  E.D.  1868.  Sixth  contribution  to  the  herpetology  of 
tropical  America.  Proc.  Acad.  Natur.  Sci.  Philadelphia 
1868:305-313. 

. 1869.  A review  of  the  species  of  the  Plethodontidae 

and  Desmognathidae.  Proc.  Acad.  Natur.  Sci.  Philadel- 
phia 21:93-118. 

Duellman,  W.E.  1963.  Amphibians  and  reptiles  of  the  rain- 
forests of  southern  El  Peten,  Guatemala.  Univ.  Kansas 
Publ.  Mus.  Natur.  Hist.  1 5(5):205-249. 

Dunn,  E.R.  1921.  Two  new  Central  American  salamanders. 
Proc.  Biol.  Soc.  Washington  34:143-145. 

. 1924.  New  salamanders  of  the  genus  Oedipus  with 

a synoptical  key.  Field  Mus.  Natur.  Hist.  Zool.  Ser.  12: 
95-100. 


Contributions  in  Science,  Number  348 


Elias:  Guatemalan  Salamanders  19 


. 1926.  The  salamanders  of  the  family  Plethodon- 

tidae.  Smith  College,  Northampton,  Mass.,  viii  + 441 

pp. 

Elias,  P.,  and  D.B.  Wake.  1983.  Nyctanolis  pernix,  a new 
genus  and  species  of  plethodontid  salamander  from 
northwestern  Guatemala  and  Chiapas,  Mexico.  Pages  1- 
12  in  A.  G.  J.  Rhodin  and  K.  Miyata  (eds.).  Advances 
in  herpetology  and  evolutionary  biology:  Essays  in  honor 
of  Ernest  E.  Williams.  Mus.  Comp.  Zool.,  Cambridge, 
Mass. 

Hanken,  J.,  J.F.  Lynch,  and  D.B.  Wake.  1980.  Salamander 
invasion  of  the  tropics.  Natur.  Hist.  89(1 2):46— 53. 

Lynch,  J.F.,  and  D.B.  Wake.  1975.  Systematics  of  the  Chi- 
ropterotriton  bromeliacea  group  (Amphibia:  Caudata), 
with  description  of  two  new  species  from  Guatemala. 
Los  Angeles  Co.  Natur.  Hist.  Mus.,  Contrib.  Sci.  265: 
1-45. 

. 1978.  A new  species  of  Chiropterotriton  (Am- 
phibia: Caudata)  from  Baja  Verapaz,  Guatemala,  with 
comments  on  relationships  among  Central  American 
members  of  the  genus.  Los  Angeles  Co.  Natur.  Hist. 
Mus.,  Contrib.  Sci.  294:1-22. 

McCoy,  C.J.,  and  C.F.  Walker.  1966.  A new  salamander 
of  the  genus  Bolitoglossa  from  Chiapas.  Occ.  Papers  Mus. 
Zool.  Univ.  Mich.  649:1-1  1. 

Rosen,  D.E.  1978.  Vicariant  patterns  and  historical  expla- 
nation in  biogeography.  Syst.  Zool.  27:159-188. 

Schmidt,  K.P.  1933.  New  reptiles  and  amphibians  from 
Honduras.  Zool.  Ser.  Field  Mus.  Nat.  Hist.  322:15-22. 

. 1936.  Guatemalan  salamanders  of  the  genus  Oe- 
dipus. Zool.  Ser.  Field  Mus.  20:135-166. 

Smith,  H.M.  1945.  Herpetological  collecting  in  banana  fields 
of  Mexico.  Ward’s  Natur.  Sci.  Bull.  1:1-7. 

Smith,  W.H.  1877.  The  tailed  amphibians.  Thesis,  Mich- 
igan Univ. 

Stuart,  L.C.  1943a.  Taxonomic  and  geographic  comments 
on  Guatemalan  salamanders  of  the  genus  Oedipus.  Misc. 
Publ.  Mus.  Zool.  Univ.  Mich.  56:1-33. 

. 1 943b.  Comments  on  the  herpetofauna  of  the  Sier- 
ra de  los  Cuchumatanes  of  Guatemala.  Occ.  Papers  Mus. 
Zool.  Univ.  Mich.  471:1-29. 

. 1948.  The  amphibians  and  reptiles  of  Alta  Verapaz, 

Guatemala.  Misc.  Publ.  Mus.  Zool.  Univ.  Mich.  69: 1 — 
109. 


. 1950.  A geographic  study  of  the  herpetofauna  of 

Alta  Verapaz,  Guatemala.  Contrib.  Lab.  Vert.  Biol.  Univ. 
Mich.  45:1-77. 

. 1952.  Some  new  amphibians  from  Guatemala.  Proc. 

Biol.  Soc.  Washington  65:1-12. 

Taylor,  E.H.  1941.  New  amphibians  from  the  Hobart  M. 
Smith  Mexican  Collections.  Univ.  Kansas  Sci.  Bull.  28: 
141-167. 

. 1944.  The  genera  of  plethodont  salamanders  in 

Mexico,  Pt.  1.  Univ.  Kans.  Sci.  Bull.  30:189-232. 

Taylor,  E.H. , and  H.M.  Smith.  1945.  Summary  of  the  col- 
lections of  amphibians  made  in  Mexico  under  the  Walter 
Rathbone  Bacon  Traveling  Scholarship.  Proc.  U.S.  Natl. 
Mus.  95:521-613. 

Wake,  D.B.,  and  A.H.  Brame,  Jr.  1963.  The  status  of  the 
plethodontid  salamander  genera  Bolitoglossa  and  Mag- 
nadigita.  Copeia  1963:382-387. 

. 1 969.  Systematics  and  evolution  of  neotropical  sal- 
amanders of  the  Bolitoglossa  helmrichi  group.  Natur. 
Hist.  Mus.  Los  Angeles  County,  Contrib.  Sci.  1 75: 1-40. 

Wake,  D.B.,  and  I.G.  Dresner.  1967.  Functional  mor- 
phology and  evolution  of  tail  autotomy  in  salamanders. 
J.  Morph.  122:265-306. 

Wake,  D.B.,  and  P.  Elias.  1983.  New  genera  and  a new 
species  of  Central  American  salamanders,  with  a review 
of  the  tropical  genera  (Amphibia,  Caudata,  Plethodon- 
tidae).  Natur.  Hist.  Mus.  Los  Angeles  County,  Contrib. 
Sci.  345:1-19. 

Wake.  D.B.,  and  J.F.  Lynch.  1976.  The  distribution,  ecol- 
ogy, and  evolutionary  history  of  plethodontid  salaman- 
ders in  tropical  America.  Bull.  Natur.  Hist.  Mus.  Los 
Angeles  Co.  25:1-65. 

. 1982.  Evolutionary  relationships  among  Central 

American  salamanders  of  the  Bolitoglossa  franklini 
group,  with  a description  of  a new  species  from  Gua- 
temala. Herpetologica  38:257-272. 

Wake,  D.B..  S.Y.  Yang,  and  T.J.  Papenfuss.  1980.  Natural 
hybridization  and  its  evolutionary  implications  in  Gua- 
temalan plethodontid  salamanders  of  the  genus  Bolito- 
glossa. Herpetologica  36:335-345. 


Submitted  6 October  1981;  accepted  23  August  1983. 


20  Contributions  in  Science,  Number  348 


Elias:  Guatemalan  Salamanders 


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SYSTEMATICS  OF  XANTUSIID  LIZARDS 
OF  THE  GENUS  LEPIDOPH  YMA 
IN  NORTHEASTERN  MEXICO 


Robert  L.  Bezy 


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SYSTEMA TICS  OF  XANTUSIID  LIZARDS 
OF  THE  GENUS  LEPIDOPHYMA 
IN  NORTHEASTERN  MEXICO 

Robert  L.  Bezy1 


ABSTRACT.  Discriminant  analyses  of  variation  among  30  scale 
characters  indicate  that  the  21  populations  of  Lepidophyma  from 
northeastern  Mexico  form  six  morphological  groups.  Two  of  27 
population  samples  from  southern  Mexico  approach  the  northern 
groups  in  discriminant  space.  When  viewed  in  terms  of  univariate 
differences  and  geographic  distribution,  the  northern  population 
groups  constitute  four  unique  morphological  entities  that  are  con- 
sidered to  represent  species  units.  The  most  distinctive  is  Lepido- 
phyma gaigeae  occurring  in  limestone  habitats  in  the  Sierra  Madre 
Oriental  of  Hidalgo  and  Queretaro.  Lepidophyma  occulor  is  known 
from  four  localities  in  the  semi-arid  Jalpan  region  of  Queretaro  and 
San  Luis  Potosi,  and  the  cavemicolous  L.  micropholis  is  confined 
to  the  Sierra  del  Abra  of  Tamaulipas  and  San  Luis  Potosi.  The  wide- 
ranging  L.  sylvaticum  includes  four  moderately  divergent  population 
groups:  northern  Madrean  (Tamaulipas  to  San  Luis  Potosi),  southern 
Madrean  (San  Luis  Potosi  to  Hidalgo),  Veracruzan,  and  western 
(Mesa  Central  of  San  Luis  Potosi  to  Nuevo  Leon). 

The  karyotypes  of  L.  gaigeae  and  L.  occulor  are  unique  within 
the  genus,  while  most  L.  sylvaticum  are  chromosomally  identical  to 
L.  micropholis.  A heteromorphism  in  microchromosomes  was  ob- 
served in  six  females  of  one  population  of  L.  sylvaticum.  and  could 
represent  either  ZW  sex  chromosomes  or  allodiploidy.  This  same 
population  plus  one  in  Queretaro  have  statistically  significantly  skewed 
sex  ratios  that  may  be  associated  with  hybridization. 

RESUMEN.  Los  analisis  discriminatorios  de  variacion  entre  30 
caracteres  de  las  escamas,  indican  que  las  2 1 poblaciones  de  Lepi- 
dophyma del  noreste  de  Mexico  forman  seis  grupos  morfologicos. 
Dos  de  los  muestreos  de  la  poblacion  del  sur  de  Mexico  se  aproximan 
a los  grupos  del  norte  en  espacio  discriminatorio.  Cuando  se  ex- 
aminan  en  terminos  de  diferencias  univariadas  y de  distribucion 
geografica,  los  grupos  de  la  poblacion  del  norte  constituyen  cuatro 
entidades  morfologicas  unicas  que  se  considera  representan  unidades 
de  especie.  La  mas  distintiva  es  Lepidophyma  gaigeae  que  vive  en 
habitats  de  piedra  caliza  en  la  Sierra  Madre  Oriental  de  Hidalgo  y 
Queretaro.  Lepidophyma  occulor  se  conoce  de  cuatro  localidades  de 
la  region  semiarida  de  Jalpan  de  Queretaro  y San  Luis  Potosi,  y el 
cavemicola  L.  micropholis  se  encuentra  confinado  a la  Sierra  del 
Abra  de  Tamaulipas  y San  Luis  Potosi.  L.  sylvaticum  de  amplia 
distribucion  en  y cerca  de  la  Sierra  Madre  Oriental  incluye  cuatro 
grupos  poblacionales  moderadamente  divergentes:  norte  (Tamau- 
lipas a San  Luis  Potosi),  sur  (San  Luis  Potosi  a Hidalgo),  Veracruz, 
y occidental  (Mesa  Central  de  San  Luis  Potosi  a Nuevo  Leon). 

Los  cariotipos  de  L.  gaigeae  y L.  occulor  son  unicos  dentro  del 
genero  mientras  que  la  mayoria  de  L.  sylvaticum  son  cromosomi- 
camente  identicos  a L.  micropholis.  Se  observo  heteromorfismo  de 
microcromosomas  en  seis  hembras  de  una  poblacion  de  L.  sylva- 
ticum y podria  representar  ya  sea  cromosomas  sexuales  ZW  o alo- 
diploidia.  Esta  misma  poblacion,  mas  una  en  Queretaro  han  torcido 
estadisticaniente  en  forma  significativa  las  proporciones  en  los  sexos 
que  pudieran  ser  asociadas  con  hibridizacion. 

INTRODUCTION 

Lizards  of  the  xantusiid  genus  Lepidophyma  range  from  Pan- 
ama to  Nuevo  Leon,  Mexico,  living  principally  in  wet  trop- 

Contributions  in  Science,  Number  349,  pp.  1-16 
Natural  History  Museum  of  Los  Angeles  County,  1984 


ical  lowland  forests  in  the  south,  but  becoming  increasingly 
restricted  to  montane  and/or  rimose  habitats  in  the  semi- 
arid  regions  to  the  north.  In  the  rugged  ranges  of  the  Sierra 
Madre  Oriental,  and  in  the  canyons  and  valleys  along  both 
of  its  flanks,  are  a morphologically  diverse  array  of  Lepi- 
dophyma populations.  While  most  of  these  populations  re- 
main taxonomically  unallocated,  four  have  been  named,  and 
two  of  these  names  have  been  alternatively  associated  with 
species  occurring  to  the  south  (e.g..  Smith,  1942;  Walker, 
1955). 

In  this  paper,  the  problems  of  discordant  morphological 
variation,  geographic  isolation,  and  small  sample  sizes  of  the 
populations  of  Lepidophyma  in  northeastern  Mexico  are  han- 
dled by  treating  each  locality  as  a separate  sample,  and  em- 
ploying multivariate  analyses  of  variation  to  identify  groups 
of  morphologically  similar  populations.  Additional  multi- 
variate comparisons  with  populations  to  the  south,  and  anal- 
yses of  univariate  differences  among  all  population  groups 
are  used  to  diagnose  morphological  species.  Names  are  then 
assigned  to  the  units  on  the  basis  of  included  type  or  topotypic 
material,  the  species  of  Lepidophyma  recognized  in  north- 
eastern Mexico  are  summarized  in  brief  accounts,  including 
comments  on  chromosomal  variation  and  skewed  sex  ratios 
in  certain  populations,  and  a key  is  presented. 

MATERIALS  AND  METHODS 

A total  of  1 52  specimens  of  Lepidophyma  from  Mexico  north 
of  19°N  latitude  were  used  in  the  analyses.  This  includes  all 
material  studied  from  the  area,  except  that  referable  to  L. 
gaigeae.  One  sample  (N  = 20)  of  the  over  500  known  spec- 
imens of  the  species  was  used  as  a reference  population.  In 
addition,  31  population  samples  (N  = 188)  from  southern 
and  western  Mexico  were  utilized  in  the  comparative  anal- 
yses. The  selection  of  19°N  latitude  as  the  southern  limit  of 
the  study  area  is  based  on  a distributional  hiatus  for  the  genus 
in  the  transvolcanic  region  (ca.  1 9-20°N),  and  on  preliminary 
observations  suggesting  that  the  populations  occurring  to  the 
north  of  this  distributional  gap  share  a number  of  unique 
morphological  similarities. 

The  localities  of  the  specimens  were  determined  on  avail- 
able maps,  and  geographic  samples  were  constituted  with  all 
specimens  from  a given  locality  (or  in  a few  instances  by 
pooling  adjacent  localities  separated  by  less  than  20  km)  to 
form  a total  of  2 1 population  samples  of  Lepidophyma  from 
northern  Mexico  (Fig.  1 ).  The  specimens  and  localities  are 
listed  in  specimens  examined,  below. 


1 . Section  of  Herpetology,  Natural  History  Museum  of  Los  An- 
geles County,  900  Exposition  Boulevard,  Los  Angeles,  California 
90007. 


ISSN  0459-8113 


The  analyses  use  a total  of  30  scale  characters,  20  meristics, 
and  1 0 ratios  of  the  relative  size  or  proportions  of  individual 
scales.  No  significant  sexual  dimorphism,  ontogenetic  vari- 
ation, or  correlation  was  detected  among  the  30  characters. 
The  characters  were  selected  largely  on  the  basis  of  their 
purported  diagnostic  strength  in  the  genus  (Bezy,  1973;  Bezy 
et  al.,  1982;  Mosauer,  1936;  Smith,  1942,  1973;  Smith  and 
Alvarez  del  Toro,  1977;  Taylor,  1939;  Walker,  1955;  Werler, 
1957;  Werler  and  Shannon,  1957). 

Scale  terminology  follows  Savage  (1963).  The  characters 
are  defined  below. 


1. 

2. 

3. 

4. 

5. 

6. 


7. 


8. 

9. 

10. 

11. 

12. 

13. 

14. 

15. 

16. 

17. 

18. 

19. 

20. 

21. 

22. 

23. 

24. 

25. 

26. 

27. 

28. 


FPT 

LTR 

DBPVR 

1WD1 

IWV1 

PTMP 


DBPVT 


GC1IL 

GUL 

PVTL 

VL 

4TL 

4TLD 

DOR 

DAPVR 

PVR 

PVS 

PVT1 

PVT2 

PVT3 

RPOL 

RPAW 

RPFML 

RPFL1 

RMW 

RNL 

RML 

RAPPSL 


Femoral  pores  (total  both  sides). 

Lateral  rows  of  tubercles  (axilla  to  groin). 
Dorsals  between  paravertebral  rows  of  tu- 
bercles. 

Dorsal  interwhorls  in  first  caudal  segment. 
Ventral  interwhorls  in  first  caudal  segment. 
Pretympanics  (total  both  sides)  separating 
postocular  from  second  postorbital  suprala- 
bial. 

Distance  between  large  paravertebral  tuber- 
cles within-row,  expressed  in  number  of  mid- 
dorsal scales. 

Gulars  contacting  first  pair  of  mfralabials. 
Gulars  (fold  to  second  infralabials). 

Large  tubercles  in  paravertebral  row  (axilla 
to  groin). 

Ventrals  (gular  to  vent;  includes  preanals). 
Fourth  toe  lamellae  (ventral). 

Fourth  toe  lamellae  divided  (i.e.,  with  ca.  mid- 
ventral  sutures). 

Dorsals  occiput  to  rump  (above  vent). 
Dorsals  in  row  immediately  above  paraver- 
tebral row  (axilla  to  groin). 

Total  scales  in  paravertebral  row  (axilla  to 
groin). 

Scales  in  paravertebral  row  (a-g)  smaller  than 
1.5  dorsals. 

Scales  in  paravertebral  row  (a-g)  larger  than 
1.5  dorsals. 

Scales  in  paravertebral  row  (a-g)  larger  than 

2.0  dorsals. 

Scales  in  paravertebral  row  (a-g)  larger  than 

3.0  dorsals. 

Length  of  postocular/length  of  orbit. 

Width  of  posterolateral  preanal/width  of  pos- 
teromedial preanal. 

Prefrontal;  length  along  midline/length  along 
lateral  border. 

Prefrontal:  length  of  mid-line  suture/length 
along  lateral  border. 

Width  of  median  (prefrontal)/anterior  width 
of  interparietal. 

Length  of  nasal/length  of  postparietal. 
Length  of  median  (prefrontal)/length  of  fron- 
tal. 

Length  (total  both  sides)  of  all  anomalous  su- 
tures on  postparietals/length  of  postparietals. 


Figure  1.  Location  of  the  21  population  samples  of  Lepidophyma 
in  Mexico  north  of  19°N.  Stippled  area  indicates  approximate  dis- 
tribution of  pine-oak  woodland  (after  Leopold,  1959).  Population 
numbers  are  those  used  throughout  the  paper  (see  Specimens  Ex- 
amined for  localities). 


29.  RPNH  Height  of  postnasal/height  of  anterior  loreal. 

30.  RSLH  Height  of  second  postorbital  supralabial/ 

height  of  first  postorbital  supralabial. 

Variation  in  the  30  characters  was  analyzed  univariately 
with  BMDP1D  for  simple  data  description,  and  multivari- 
ately  with  BMDP7M  for  stepwise  discriminant  analysis  (Dix- 
on, 1981).  In  all  discriminant  analyses  the  a priori  groups 
were  individual  population  samples  rather  than  population 
groups  or  species. 

RESULTS  AND  DISCUSSION 

In  the  following  sections  the  results  of  discriminant  analyses 
of  populations  of  Lepidophyma  from  northern  Mexico  are 
used  to  identify  northern  population  groups,  which  in  turn 
are  compared  with  populations  from  southern  Mexico.  The 
northern  population  groups  are  then  viewed  relative  to  their 
univariate  differences  and  geographic  relationships  to  arrive 
at  the  definition  of  unique  morphological  units.  Finally,  names 
are  allocated  to  these  units  (morphospecies)  on  the  basis  of 
included  topotypic  and/or  type  material,  and  each  species  is 
briefly  summarized. 

NORTHERN  MEXICO  POPULATIONS 

The  initial  discriminant  analysis  utilized  30  characters  and 
18  of  the  21  populations  from  northeastern  Mexico  (Fig.  1, 


2 Contributions  in  Science,  Number  349 


Bezy:  Systematics  of  Lepidophyma  in  Mexico 


_J I I I I — 

-14  -7  0 7 14 


Figure  2.  Nineteen  population  samples  of  Lepidophyma  from 
northeastern  Mexico  plotted  on  the  first  two  canonical  variables  for 
25  characters.  Population  centroids  are  indicated  by  dots,  and  the 
number  of  the  sample  is  placed  along  the  line  enclosing  all  included 
specimens  (lower  case  letters).  Upper  case  letters  indicate  population 
groups  identified  by  the  analysis. 


Table  1).  In  samples  3,  5,  and  19  there  are  no  individuals 
on  which  all  of  the  characters  could  be  scored.  The  analysis 
resulted  in  high  resolution  of  the  populations  in  that  99  per- 
cent (127/128)  of  the  individuals  were  “correctly”  assigned 
by  the  posterior  classification  to  the  locality  sample  of  which 
they  were  a member  (one  specimen  of  sample  7 was  mis- 
assigned  to  6).  A second  analysis  was  performed  excluding 
five  characters  (IWD1,  IWV1,  RPFML,  RPFL1,  RML)  in 
order  to  allow  inclusion  of  sample  5 (Fig.  2).  Samples  3 and 
19  were  not  included  in  any  of  the  multivariate  analyses  due 
to  the  limitations  of  the  data  available  from  them.  While 
both  analyses  produced  similar  results,  the  reduction  in  char- 
acters of  the  second  lowered  the  accuracy  of  the  posterior 
classification  (96%;  126/131). 

From  the  second  analysis,  eight  population  groups  were 
identified  on  the  basis  of  overlap  or  juxtaposition  of  the 
included  samples  and  the  distance  between  groups  in  dis- 
criminant space  (Fig.  2).  The  first  canonical  variable  accounts 
for  57  percent  of  the  variation,  is  most  heavily  loaded  with 
LTR.  PVS,  PVR,  DOR,  and  GUL  (in  order  of  decreasing 
weight),  and  places  group  A at  one  end,  and  B,  C,  and  D at 
the  other,  with  E,  F,  G,  and  H occupying  intermediate  po- 
sitions. The  second  coordinate  has  heavy  loadings  for  PVS, 
PVR,  PVT1,  LTR,  and  FPT,  accounts  for  21  percent  of  the 
variation,  and  effectively  separates  groups  E,  G,  and  H from 
one  another. 

The  sample  comprising  group  A (21)  is  highly  isolated  in 
discriminant  space  from  all  other  populations,  suggesting  it 
is  not  a member  of  the  same  morphological  complex.  Group 
D is  a discrete  cluster  of  four  overlapping  populations  (13- 
lb)  that  is  approximately  equidistant  from  B (17,  18)  and  C 
(20).  The  three  populations  of  group  E (9-1 1)  form  a mod- 
erately tight  cluster  that  is  only  weakly  separated  from  the 
loosely  associated  populations  of  G (5-8)  and  the  one  spec- 
imen (12)  comprising  the  intermediate  group  F.  The  three 
populations  of  group  H ( 1 , 2,  4)  are  well  separated  from  their 
nearest  discriminant  neighbor,  group  G. 


(J I I I l_ 

-14  -7  0 7 14 


Figure  3.  Twelve  population  groups  of  Lepidophyma  from  Mexico 
plotted  on  the  first  two  canonical  variables  for  30  characters.  Lines 
enclose  all  individuals  comprising  each  of  the  eight  northern  (A-G) 
and  four  southern  (I-L)  population  groups. 


COMPARISONS  WITH  SOUTHERN  GROUPS 

Twenty-seven  samples  from  southern  Mexico  were  com- 
pared with  the  18  northern  populations  to  identify  those  that 
might  be  closest  morphologically  to  northern  groups.  The 
initial  discriminant  anlaysis  utilized  30  characters  and  a total 
of  288  specimens  arrayed  in  45  populations  (Fig.  3),  and 
produced  high  resolution  of  the  populations  in  that  the  ac- 
curacy of  the  posterior  classification  was  98  percent.  The  first 
canonical  variable  is  most  heavily  loaded  with  PVS,  DBPVT, 
LTR,  GUL,  and  PVR,  and  expresses  43  percent  of  the  total 
dispersion;  the  second  expresses  2 1 percent  and  is  dominated 
by  LTR,  PVS,  PVTL,  FPT,  and  RPAW.  The  graph  (Fig.  3) 
was  used  primarily  to  identify  those  southern  population 
groups  that  are  multivariately  most  similar  to  the  northern 
ones  and  which  are  further  resolved  in  subsequent  analyses 
containing  fewer  populations. 

All  but  one  (Fj  of  the  eight  northern  groups  identified 
in  the  previous  analysis  remain  separated  from  each  other, 
although  they  are  approached  or  overlapped  by  three  south- 
ern groups  (J,  K,  L)  (Fig.  3).  Northern  groups  B,  C,  and  D 
were  well  separated  from  both  southern  and  northern  pop- 
ulations and  thus  are  not  included  in  the  subsequent  analyses. 
Southern  group  I is  also  strongly  separated  from  all  popu- 
lations, and  its  nearest  discriminant  neighbor  is  another 
southern  group  (J).  Consequently,  it  was  also  excluded  from 
further  analysis.  In  the  following  analyses,  northern  groups 
E through  H are  compared  in  greater  detail  first  with  K and 
L,  and  then  with  J. 

The  10  populations  of  northern  groups  E,  F,  G,  and  H 
were  analyzed  together  with  the  four  populations  of  southern 
groups  K and  L (Fig.  4).  The  posterior  classification  was  98 
percent  ( 1 2 1/123)  accurate,  one  specimen  of  sample  11  being 
misassigned  to  10  (both  group  E)  and  one  of  sample  43  to 
44  (both  group  K).  The  first  canonical  variable  accounts  for 
60  percent  of  the  total  dispersion,  is  heavily  loaded  with 


Contributions  in  Science,  Number  349 


Bezy:  Systematics  of  Lepidophyma  in  Mexico  3 


Table  1.  Variation  among  30  scale  characters  for  21  population  samples  of  Lepidophyma  from  northern  Mexico.  Sample  size  is  (in  parentheses) 
under  each  of  the  population  numbers  (POP).  In  each  cell  the  upper  number  is  the  mean;  the  middle,  the  standard  error;  and  the  lower,  the 
range.  See  text  for  character  abbreviations  and  locality  data. 


POP 

FPT 

LTR 

DBPVR 

1 WD1 

IWV1 

PTMP 

DBPVT 

GC1IL 

GUL 

PVTL 

VL 

4TL 

4TLD 

DOR 

DAPVR 

PVR 

1 

31.3 

18.1 

3.17 

3.0 

2.1 

2.1 

2.83 

.3 

45.0 

15.9 

36.1 

25.3 

8.1 

161.2 

83.2 

44.8 

(15) 

.37 

.59 

.093 

0 

.09 

.07 

.080 

.13 

.59 

.21 

.15 

.33 

.56 

1.88 

.99 

1.13 

29-34 

15-22 

2. 5-4.0 

3 

2-3 

2-3 

2. 5-3. 5 

0-1 

42-49 

15-17 

35-37 

24-29 

4-12 

150-174 

77-91 

39-53 

2 

29.0 

19.0 

2.50 

3.0 

2.0 

3.0 

2.50 

1.0 

46.0 

17.0 

35.0 

26.0 

7.0 

164.0 

83.0 

43.0 

u) 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

29 

19 

2.5 

3 

2 

3 

2.5 

1 

46 

17 

35 

26 

7 

164 

83 

43 

3 

29.0 

- 

3.50 

2.0 

2.0 

2.0 

3.00 

1.0 

46.0 

- 

- 

20.0 

5.0 

- 

- 

- 

U) 

0 

- 

0 

0 

0 

0 

0 

0 

0 

— 

- 

0 

0 

- 

- 

- 

29 

- 

3.5 

2 

2 

2 

3.0 

1 

46 

- 

- 

20 

5 

- 

- 

- 

4 

27.0 

17.0 

3.38 

2.5 

1.5 

2.3 

3.00 

.3 

45.0 

16.0 

35.3 

23.0 

6.3 

172.5 

91.0 

47.0 

(4) 

.41 

0 

.125 

.58 

.29 

.25 

0 

.25 

.71 

.41 

.25 

.41 

.25 

3.43 

1.47 

1.08 

26-28 

17 

3.0-3. 5 

2-3 

1-2 

2-3 

3.0 

0-1 

43-46 

15-17 

35-36 

22-24 

6-7 

163-178 

87-94 

44-49 

5 

28.0 

31.0 

5.00 

- 

- 

1.0 

4.50 

3.0 

56.0 

15.0 

38.0 

31.0 

15.0 

207.0 

98.0 

63.0 

U) 

0 

0 

0 

- 

- 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

28 

31 

5.0 

- 

- 

1 

4.5 

3 

56 

15 

38 

31 

15 

207 

98 

63 

6 

27.8 

29.3 

3.89 

3.2 

1.8 

2.0 

3.08 

.5 

48.3 

16.8 

35.8 

26.3 

14.6 

182.1 

93.5 

51.0 

(.32) 

.45 

.40 

.122 

.52 

.11 

.15 

.106 

.11 

.49 

.35 

.23 

.33 

.53 

1.96 

1.26 

.87 

24-35 

24-34 

3. 0-5.0 

2-4 

0-3 

0-4 

1. 0-4.0 

0-2 

43-55 

15-23 

34-39 

23-30 

9-23 

166-217 

83-1 12 

42-62 

7 

28.0 

29.0 

4.83 

3.3 

2.0 

2.0 

3.50 

.3 

51.0 

16.7 

35.3 

25.3 

13.3 

206.3 

107.0 

57.0 

(4) 

1.53 

1.00 

.167 

.58 

0 

0 

.289 

.25 

1.78 

.33 

.88 

.67 

.88 

5.36 

5.51 

5.57 

26-31 

27-30 

4. 5-5.0 

3-4 

2 

2 

3. 0-4.0 

0-1 

48-55 

17-18 

34-37 

24-26 

12-15 

196-214 

96-113 

46-64 

8 

26.2 

27.8 

4.08 

3.5 

2.5 

2.2 

3.08 

1.0 

45.5 

18.0 

36.0 

27.7 

14.2 

191.3 

97.8 

53.8 

(6) 

.40 

1.14 

.201 

.55 

.22 

.31 

.201 

0 

1.09 

1.83 

.37 

.56 

.98 

2.91 

2.12 

3.28 

25-27 

25-31 

3. 5-5.0 

3-4 

2-3 

1-3 

2. 5-4.0 

1 

43-50 

15-27 

35-37 

26-30 

1 1-18 

180-199 

91-104 

42-64 

9 

26.0 

36.0 

4.00 

3.0 

2.0 

2.0 

2.50 

0 

42.0 

27.0 

37.0 

24.0 

14.0 

191.0 

89.0 

56.0 

(!) 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

26 

36 

4.0 

3 

2 

2 

2.5 

0 

42 

27 

37 

24 

14 

191 

89 

56 

10 

26.1 

33.7 

4.64 

3.1 

1.9 

1.6 

1.71 

.6 

44.0 

24.9 

34.8 

25.5 

17.7 

180.1 

91.0 

52.2 

(22) 

.31 

.44 

.082 

.47 

.06 

.15 

.146 

.11 

.42 

1.47 

.24 

.14 

.52 

1.33 

1.25 

1.18 

24-30 

31-38 

4. 0-5.0 

2-4 

1-2 

0-2 

1. 0-3.0 

0-1 

41-49 

15-39 

33-37 

24-27 

13-23 

165-188 

78-99 

40-66 

1 1 

25.2 

34.2 

4.25 

4.0 

2.2 

2.2 

3.25 

1.1 

47.8 

19.0 

36.6 

25.7 

18.8 

184.6 

95.5 

61.4 

(10) 

.55 

.51 

112 

.67 

.20 

.13 

.186 

.18 

.83 

1.09 

.16 

.45 

.57 

2.57 

1.86 

1.63 

23-28 

32-36 

3. 5-4.5 

3-5 

1-3 

2-3 

2. 5-4.0 

0-2 

45-54 

15-25 

36-37 

23-28 

16-21 

167-192 

88-104 

56-73 

12 

31.0 

35.0 

5.00 

4.0 

2.0 

3.0 

4.00 

0 

47.0 

17.0 

36.0 

28.0 

18.0 

205.0 

103.0 

53.0 

u) 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

31 

35 

5.0 

4 

2 

3 

4.0 

0 

47 

17 

36 

28 

18 

205 

103 

53 

13 

32.7 

29.3 

5.67 

5.0 

3.0 

.7 

4.67 

.3 

62.0 

16.0 

36.3 

25.3 

12.3 

243.3 

126.7 

70.3 

(3) 

1.45 

1.20 

.167 

0 

0 

.67 

.333 

.33 

3.21 

.58 

.33 

.88 

1.20 

4.41 

7.26 

4.81 

30-35 

27-31 

5. 5-6.0 

5 

3 

0-2 

4. 0-5.0 

0-1 

57-68 

15-17 

36-37 

24-27 

10-14 

235-250 

115-140 

61-77 

14 

32.6 

30.6 

6.27 

4.7 

2.5 

1.2 

4.68 

.4 

62.4 

16.2 

35.9 

26.5 

13.9 

244.2 

122.5 

73.4 

(//) 

.62 

.59 

.124 

.47 

.16 

.25 

.122 

.16 

.88 

.23 

.21 

.43 

.64 

1.50 

2.14 

2.00 

29-36 

28-35 

6. 0-7.0 

4-5 

2-3 

0-2 

4. 0-5.0 

0-1 

58-68 

15-17 

35-37 

24-29 

1 1-17 

235-251 

112-134 

65-85 

15 

29.5 

29.2 

4.92 

3.7 

1.8 

1.8 

5.00 

.5 

56.8 

15.8 

36.3 

26.0 

13.2 

237.5 

128.0 

62.5 

(6) 

.56 

.40 

.239 

.52 

.17 

.17 

0 

.34 

.70 

.31 

.21 

.45 

.65 

2.43 

2.36 

2.14 

28-3 1 

28-31 

4.0-5. 5 

3-4 

1-2 

1-2 

5.0 

0-2 

55-60 

15-17 

36-37 

25-28 

1 1-15 

231-245 

1 19-135 

52-66 

16 

29.8 

30.2 

4.80 

4.0 

2.2 

2.0 

4.90 

.8 

58.6 

15.6 

36.0 

27.0 

16.8 

239.4 

120.4 

60.8 

(5) 

.80 

.49 

.123 

0 

.20 

0 

.100 

.20 

.81 

.40 

.32 

.71 

.66 

1.40 

2.42 

2.43 

28-32 

29-31 

4. 5-5.0 

4 

2-3 

2 

4. 5-5.0 

0-1 

56-61 

15-17 

35-37 

25-29 

15-19 

236-244 

112-127 

56-70 

17 

19.0 

22.5 

5.00 

3.5 

2.0 

2.0 

4.00 

0 

67.0 

17.0 

37.5 

25.0 

6.0 

228.5 

113.0 

54.0 

(3) 

1.00 

1.50 

0 

.71 

0 

0 

0 

0 

4.00 

0 

.50 

0 

0 

4.50 

1.00 

1.00 

18-20 

21-24 

5.0 

3-4 

2 

2 

4.0 

0 

63-71 

17 

37-38 

25 

6 

224-233 

112-114 

53-55 

18 

19.0 

21.3 

4.67 

3.0 

1.7 

2.0 

4.00 

0 

61.0 

15.7 

35.3 

24.7 

7.3 

239.3 

1 18.3 

60.7 

(3) 

1.15 

.67 

.441 

0 

.33 

0 

0 

0 

.58 

.67 

.33 

.33 

.88 

1.33 

1.86 

4.63 

17-21 

20-22 

4.0-5. 5 

3 

1-2 

2 

4.0 

0 

60-62 

15-17 

35-36 

24-25 

6-9 

238-242 

1 16-122 

53-69 

19 

19.5 

24.0 

5.00 

4.0 

2.0 

2.0 

4.50 

— 

66.0 

17.0 

36.5 

24.0 

8.5 

- 

- 

- 

(2) 

.50 

0 

0 

0 

0 

0 

.500 

- 

1.00 

1.00 

.50 

0 

.50 

- 

- 

- 

19-20 

24 

5.0 

4 

2 

2 

4. 0-5.0 

- 

65-67 

16-18 

36-37 

24 

8-9 

- 

- 

- 

20 

19.5 

21.5 

3.00 

3.0 

2.0 

2.5 

4.00 

0 

60.0 

15.5 

35.5 

22.0 

4.5 

216.5 

101.0 

53.5 

(2) 

1.50 

1.50 

0 

0 

0 

.50 

0 

0 

1.00 

.50 

.50 

0 

1.50 

3.50 

1.00 

4.50 

18-21 

20-23 

3.0 

3 

2 

2-3 

4.0 

0 

59-61 

15-16 

35-36 

22 

3-6 

213-220 

100-102 

49-58 

21 

33.5 

46.5 

4.02 

2.2 

2.0 

3.8 

2.54 

.5 

36.4 

11.3 

34.3 

26.3 

11.1 

133.5 

64.2 

51.2 

(20) 

.28 

.54 

.057 

.38 

0 

.24 

.098 

.11 

.41 

.73 

.19 

.32 

.46 

1.09 

.69 

1.08 

32-37 

43-50 

3. 5-5.0 

2-3 

2 

2-6 

2. 0-3.0 

0-1 

33-39 

6-18 

33-36 

25-30 

7-16 

126-142 

59-68 

44-58 

4 Contributions  in  Science,  Number  349 


Bezy:  Systematics  of  Lepidophyma  in  Mexico 


Table  1.  Continued. 


POP 

PVS 

PVT  1 

PVT2 

PVT3 

RPOL 

RPAW 

RPFML 

RPFL1 

RMW 

RNL 

RML 

RAPPSL 

RPNH 

RSLH 

1 

13.9 

30.0 

20.2 

15.7 

.229 

.603 

.459 

.382 

.117 

.136 

.100 

0 

.704 

1.079 

(15) 

.98 

.55 

.81 

.21 

.0249 

.0311 

.0409 

.0475 

.0514 

.0086 

.0491 

0 

.0118 

.0476 

7-20 

26-34 

15-28 

15-17 

. 1 5-56 

.39-82 

.1 1-.70 

.1 1-.65 

0-.56 

o 

O' 

o 

0-.63 

0 

.65-8 1 

.91-1.61 

2 

15.0 

28.0 

17.0 

36.0 

.247 

.788 

.456 

.456 

0 

.121 

0 

0 

.754 

.890 

(/) 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

15 

28 

17 

16 

.25 

.79 

.46 

.46 

0 

.12 

0 

0 

.75 

.89 

3 

— 

— 

- 

- 

.124 

- 

.401 

.351 

0 

.309 

0 

0 

.723 

1.245 

(/> 

- 

- 

- 

— 

0 

— 

0 

0 

0 

0 

0 

0 

0 

0 

- 

- 

- 

- 

.12 

- 

.40 

.35 

0 

.31 

0 

0 

.72 

1.25 

4 

15.0 

32.0 

26.5 

16.3 

.181 

.706 

.485 

.485 

0 

.238 

0 

.221 

.907 

1.099 

(4) 

1.47 

1.08 

.87 

.48 

.0184 

.0289 

.0321 

.0321 

0 

.0151 

0 

.2214 

.1605 

.0560 

12-19 

29-34 

24-28 

15-17 

. 1 4-23 

,64-.77 

,42-,54 

.42-.  54 

0 

,22-.28 

0 

0-.89 

.73-1.39 

.98-1.21 

5 

43.0 

20.0 

15.0 

14.0 

.241 

.400 

- 

- 

0 

.072 

- 

0 

.742 

1.156 

(1) 

0 

0 

0 

0 

0 

0 

- 

- 

0 

0 

- 

0 

0 

0 

43 

20 

15 

34 

.24 

.40 

- 

- 

0 

.07 

- 

0 

.74 

1.16 

6 

22.1 

28.9 

21.4 

12.0 

.181 

.539 

.564 

.357 

.272 

.171 

.375 

.007 

.781 

1.172 

(32) 

1.25 

.72 

.84 

.59 

.0075 

.0132 

.0222 

.0362 

.0612 

.0082 

.0865 

.0042 

.0236 

.0218 

12-37 

20-35 

15-34 

4-18 

.1 1-.27 

41-.69 

.38-.S4 

0-.71 

0-.84 

. 1 1— .29 

0-1.26 

0-.1  1 

.66-1.40 

.86-1.41 

7 

26.3 

30.7 

21.3 

15.3 

.158 

.593 

.623 

.522 

0 

.133 

0 

0 

.783 

1.279 

(4) 

6.23 

.67 

2.85 

.33 

.0006 

.0382 

.0177 

.0203 

0 

.0318 

0 

0 

.0121 

.0246 

14-34 

30-32 

18-27 

15-16 

.16 

.53-.66 

.60-.66 

.48-. 54 

0 

.08-.  19 

0 

0 

o 

00 

f 

SO 

r- 

1.23-1.32 

8 

27.5 

26.3 

19.3 

14.0 

.181 

.545 

.513 

.344 

.213 

.164 

.319 

.46 

.727 

1.113 

(6) 

4.43 

1.65 

2.01 

1.03 

.0145 

.0354 

.0504 

.0800 

.1265 

.0130 

.2134 

.3021 

.0282 

.0359 

12-38 

19-30 

16-29 

9-16 

.13-.23 

.40-.66 

.36-68 

0-.52 

0— .77 

. 1 3-22 

0-1.32 

0-1.65 

.64-.81 

1.01-1.23 

9 

24.0 

32.0 

27.0 

0 

.154 

.300 

.754 

.422 

.623 

.132 

.682 

0 

.787 

1.253 

(/) 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

24 

32 

27 

0 

.15 

.30 

.75 

.42 

.62 

.13 

.68 

0 

.79 

1.25 

10 

21.0 

31.2 

24.0 

2.5 

.188 

.497 

.540 

.455 

.141 

.167 

.109 

.017 

.856 

1.180 

(22) 

1.67 

.94 

1.02 

.73 

.0140 

.0190 

.0218 

.0259 

.0585 

.01 10 

.0483 

.0173 

.0283 

.0338 

7-42 

22-37 

17-33 

0-13 

.07-32 

.31-68 

.35-.82 

. 1 4—65 

0-.95 

.09-27 

0-.85 

0-.38 

.71-1.37 

.95-1.55 

1 1 

31.5 

29.9 

19.1 

2.9 

.176 

.589 

.475 

.480 

.045 

.172 

.022 

.140 

.803 

1.141 

(10) 

1.89 

1.04 

.99 

.75 

.01 19 

.0364 

.0322 

.0433 

.0284 

.0160 

.0137 

.1102 

.0209 

.0369 

21-40 

26-35 

16-25 

0-6 

. 1 0— .22 

.39-. 75 

.27-.69 

.27-.81 

0-.28 

.12-28 

0-.13 

0-1.12 

.71-9 1 

.84-1.25 

12 

20.0 

33.0 

22.0 

9.0 

.200 

.466 

1.071 

.283 

.729 

.106 

.739 

0 

.750 

1.141 

(1) 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

20 

33 

22 

9 

.20 

.47 

1.07 

.28 

.73 

.11 

.74 

0 

.75 

1.14 

13 

42.0 

28.3 

23.3 

15.0 

.146 

.498 

.562 

.611 

0 

.188 

0 

0 

.744 

1.168 

(3) 

5.13 

2.85 

3.84 

1.15 

.0042 

.1159 

.0405 

.0654 

0 

.0051 

0 

0 

.0342 

.1171 

35-52 

25-34 

19-31 

13-17 

.14-.  15 

.27-63 

.49-63 

.49-.  71 

0 

. 1 8-.  1 9 

0 

0 

.70-8 1 

.94-1.32 

14 

42.6 

30.7 

23.5 

15.6 

.168 

.570 

.579 

.579 

0 

.129 

0 

0 

.745 

1.176 

an 

2.38 

.76 

1.26 

.34 

.0062 

.0313 

.0321 

.0321 

0 

.0027 

0 

0 

.0276 

.0701 

32-56 

26-34 

17-29 

14-18 

. 14— .20 

,35-,74 

.46-.  76 

.46-. 76 

0 

.11-.  15 

0 

0 

.66-.95 

.95-1.76 

15 

26.7 

35.8 

30.3 

17.7 

.140 

.728 

.528 

.528 

0 

.181 

0 

0 

.754 

1.11  1 

(6) 

2.20 

1.22 

1.23 

.67 

.0085 

.0584 

.0198 

.0198 

0 

.0129 

0 

0 

.0184 

.0427 

17-32 

33-40 

26-35 

16-20 

12-.18 

46-.85 

.48-. 62 

.48-. 62 

0 

.14-22 

0 

0 

.70-.83 

.96-1.21 

16 

29.8 

31.0 

28.2 

16.2 

.189 

.500 

.544 

.544 

0 

.129 

0 

0 

.743 

1.114 

(5) 

2.63 

.45 

.80 

.58 

.0177 

.0185 

.0468 

.0468 

0 

.0123 

0 

0 

.0236 

.0259 

25-40 

30-32 

26-30 

15-18 

. 14-.24 

,44-.54 

.42-.68 

.42-.68 

0 

.10-15 

0 

0 

.69-8 1 

1.02-1.17 

17 

26.5 

27.5 

31.5 

19.5 

.151 

.537 

.670 

.670 

0 

.136 

0 

0 

.677 

.954 

(2) 

5.50 

4.50 

1.50 

1.50 

.0217 

.0020 

.0049 

.0049 

0 

.0232 

0 

0 

.0636 

.0087 

21-32 

23-32 

30-33 

18-21 

. 1 3-.  1 7 

.54 

.67 

.67 

0 

.1 1-.16 

0 

0 

.61-74 

,95-,96 

18 

30.3 

30.3 

30.3 

20.7 

.181 

.566 

.747 

.747 

0 

.156 

0 

.037 

.775 

.819 

(3) 

4.91 

.33 

.33 

1.45 

.0206 

.0496 

.0247 

.0247 

0 

.0102 

0 

.0370 

.0039 

.0343 

19 
(2) 

20 

22-39 

30-31 

30-31 

18-23 

. 1 5-,22 

.178 

.0018 

.18 

.145 

.55-.66 

.70-.77 

.70-.78 

0 

.14-.  18 

0 

0-.1 1 

.77-78 

.76-.88 

22.0 

31.5 

30.0 

16.0 

.629 

.709 

.709 

0 

.090 

0 

0 

.776 

.805 

(2) 

4.00 

.50 

0 

0 

.0251 

.0592 

.0443 

.0443 

0 

.0059 

0 

0 

.0281 

.0216 

18-26 

31-32 

30 

16 

. 1 2-.  1 7 

,57-.69 

,67-,75 

,67-,75 

0 

.08-10 

0 

0 

.75— .8 1 

,78-,83 

21 

36.5 

14.6 

3.9 

0 

.208 

.407 

.491 

.405 

.094 

.199 

.137 

.052 

.744 

.745 

(20) 

1.97 

1.05 

.74 

0 

.0056 

.0168 

.0264 

.0367 

.0400 

.0076 

.0587 

.0323 

.0087 

.0155 

22-50 

8-22 

0-1  1 

0 

. 1 5-26 

.27-.54 

.28-.  71 

0-.63 

0-.62 

.15-.  26 

0-.84 

0-.59 

O' 

00 

1 

bo 

O 

.59-.  86 

Contributions  in  Science,  Number  349 


Bezy:  Systematics  of  Lepidophyma  in  Mexico  5 


Figure  4.  Fourteen  samples  of  Lepidophyma  of  two  southern  (K. 
and  L)  and  four  northern  (E-H)  population  groups  plotted  on  the 
first  two  canonical  variables  for  30  characters.  Dots  are  centroids; 
sample  numbers  are  along  lines  enclosing  all  included  specimens. 

4TLD,  LTR,  PVS,  PVR,  and  RMW;  the  second  explains  16 
percent  of  the  variation  and  is  most  weighted  with  PVS,  PVR, 
PVT  1 , FPT,  and  PVT2.  Southern  groups  K and  L are  strongly 
separated  from  H,  their  nearest  discriminant  neighbor  among 
northern  groups.  However,  one  population  of  group  H (4)  is 
separated  from  the  other  two  populations  of  the  group  in  the 
direction  of  group  L.  The  relationships  of  population  4 are 
discussed  further  on  p.  7. 

The  1 7 populations  of  group  J were  analyzed  together  with 
northern  groups  E through  H (Fig.  5).  The  accuracy  of  the 
posterior  classification  was  98  percent  (128/131),  two  spec- 
imens of  sample  1 I being  misclassified  as  10  (both  group  E). 
The  first  variable  accounts  for  42  percent  of  the  total  dis- 
persion and  is  influenced  most  by  PVS,  PVR,  PVT1,  FPT, 
and  LTR;  the  second  expresses  26  percent  and  has  heavy 
loadings  for  PVS,  PVT1,  PVR,  LTR,  and  DAPVR.  The  five 
groups  are  separated  from  one  another,  although  one  pop- 
ulation of  group  J (27)  is  separated  from  the  remainder  of 
the  group  and  is  placed  intermediate  between  J and  E and 
F,  and  two  individuals  of  sample  37  (group  J)  approach 
group  G. 

GROUP  ANALYSES 

The  multivariate  relationships  of  the  46  populations  de- 
scribed above  are  here  considered  in  respect  to  univariate 
similarities  or  differences  between  population  groups  (Table 
2)  and  to  geographic  distributions  (Fig.  6)  in  order  to  arrive 
at  the  definition  of  morphologically  diagnosable  units  of  Lep- 
idophyma occurring  in  northern  Mexico.  It  is  anticipated  that 
the  resultant  units  should  consist  of  groups  of  populations 
that  are  overlapping  orjuxtaposed  in  discriminant  space,  that 
can  be  diagnosed  by  one  or  more  univariate  characters,  that 
are  not  linked  to  other  groups  by  univariately,  multivariately, 
and  geographically  intermediate  populations,  and  that  thus 
represent  morphospecies.  For  a genus  such  as  Lepidophyma, 
in  which  sympatry  is  rare,  discordant  variation  common, 
and  populations  often  disjunct  and  represented  by  small  sam- 
ple sizes,  such  morphologically  defined  units  are  initial  species 
hypotheses  to  be  tested  by  securing  additional  samples  and 
information  (e.g.,  allozyme  data). 

Groups  E,  F,  and  G are  positioned  nearest  each  other  in 
the  four  discriminant  analyses  (Figs.  2-5)  and  they  overlap 


Figure  5.  Twenty-seven  samples  of  Lepidophyma  of  one  southern 
(.1)  and  four  northern  (E-H)  population  groups  plotted  on  the  first 
two  canonical  variables  for  30  characters.  Presentation  as  in  Fig.  4. 

in  all  individual  characters  (Table  2).  The  three  appear  to 
represent  a single  species  unit  EFG  distributed  along  the 
Sierra  Madre  Oriental  from  southern  Tamaulipas  to  Vera- 
cruz (Fig.  6). 

The  nearest  geographic  and  discriminant  neighbor  of  group 
H among  northern  populations  is  group  G (Figs.  2 and  6), 
from  which  it  differs  (=no  overlap  in  range  of  variation)  in 
LTR  (Table  2).  The  decision  as  to  whether  H should  be 
considered  specifically  distinct  from  EFG  is  complicated  by 


Figure  6.  Distribution  of  eight  population  groups  of  Lepidophyma 
in  northeastern  Mexico.  Lines  enclose  the  samples  (numbers)  in- 
cluded in  the  groups  (letters). 


6 Contributions  in  Science,  Number  349 


Bezy:  Systematics  of  Lepidophyma  in  Mexico 


Figure  7.  Lateral  body  surface  of  specimens  of  Lepidophyma  of 
groups  A (upper,  sample  21,  AMNH  13879)  and  E (middle,  sample 
1 1,  LACM  106742;  and  lower,  sample  10,  LACM  109771). 


Figure  8.  Lateral  body  surface  of  specimens  of  Lepidophyma  of 
groups  G (upper,  sample  6,  UMMZ  1 02980;  middle,  sample  8,  LACM 
131 145)  and  H (lower,  sample  2,  EAL  4644). 


the  presence  of  intermediate  states,  observed  in  the  lateral 
tubercle  rows,  that  are  not  expressed  in  the  LTR  counts.  In 
some  specimens,  the  low  number  of  lateral  tubercle  rows  that 
characterizes  group  H results  from  a slight  reduction  of  some 
of  the  rows  in  terms  of  the  distance  they  extend  above  the 
ventrals  and  the  relative  size  of  the  tubercles  which  compose 
them  (Figs.  7-8).  While  uniform  criteria  were  employed 
throughout  the  study  to  determine  which  rows  to  include  in 
the  counts,  for  some  of  the  specimens  in  groups  G and  H 
the  decision  was  difficult  and  repeatability  of  the  counts  was 
low.  The  difference  in  LTR  between  H and  EFG  is  thus  less 
discrete  than  suggested  by  the  counts  and  is  bridged  by  in- 
termediate morphological  states.  While  additional  specimens 
and  information  (e.g.,  allozyme  data)  are  needed  to  fully 
evaluate  this  situation,  it  seems  best  not  to  place  emphasis 
on  the  differences  in  LTR  number,  and  to  recognize  a single 
species  unit  composed  of  groups  E,  F,  G,  and  H. 

In  one  of  the  analyses,  population  4 is  slightly  separated 
from  the  other  two  populations  of  group  H (1,  2)  in  the 
direction  of  southern  group  L from  coastal  Michoacan  (Fig. 
4).  For  a number  of  characters  (e.g.,  LTR,  IWD 1 , GUL,  4TL, 
and  4TLD),  the  mean  for  population  4 is  intermediate  be- 
tween L and  the  Nuevo  Leon  populations  (1,2)  (Tables  1 


and  2).  Nevertheless,  population  4 differs  from  L in  four 
characters  (FPT,  PTMP,  4TLD,  DOR),  and  from  the  Nuevo 
Leon  populations  (1,  2)  in  one  (FPT).  Evaluation  of  these 
differences  is  hampered  by  the  small  sample  sizes  of  popu- 
lation 4 (N  = 4)  and  group  L (N  = 3).  To  estimate  the  range 
of  variation  of  4 and  L that  would  be  expected  with  larger 
sample  sizes,  three  standard  deviations  of  population  1 (N  = 

1 5)  were  added  to  and  subtracted  from  the  means  of  each  of 
the  characters  to  encompass  99.7  percent  of  the  population 
(Simpson,  Roe,  and  Lewontin,  1 960;  1 39).  The  estimated  range 
of  population  4 overlaps  the  observed  range  of  population  1 
in  all  characters,  but  is  separated  from  the  estimated  range 
of  group  L in  FPT  (23-31  vs.  13-21)  and  PTMP  (1. 5-3.0 
vs.  3. 2-4. 8).  While  additional  material  is  necessary  to  fully 
evaluate  the  relationships  of  population  4,  the  information 
at  hand  suggests  that  it  should  be  considered  a member  of 
group  EFGH.  Further  collecting  along  the  western  flank  of 
the  Sierra  Madre  Oriental  seems  likely  to  produce  material 
linking  the  Sierra  Alvarez  population  (4)  geographically  and 
morphologically  with  the  Nuevo  Leon  populations  (1,2)  (Fig. 
6). 

Two  populations  of  southern  group  J (27,  37)  approach 
EFGH  in  discriminant  space  (Fig.  5).  The  two  groups  differ 


Contributions  in  Science,  Number  349 


Bezy:  Systematics  of  Lepidophyma  in  Mexico  7 


Table  2.  Variation  among  30  scale  characters  for  eight  population  groups  of  Lepidophyma  in  northern  Mexico.  Presentation  as  in  Table  1. 


Group 

FPT 

LTR 

DBPVR 

IWD1 

IWV1 

PTMP 

DBPVT 

GC1IL 

GUL 

PVTL 

VL 

4TL 

4TLD 

DOR 

DAPVR 

PVR 

A 

33.5 

46.5 

4.03 

2.2 

2.0 

3.8 

2.54 

.5 

36.4 

1 1.3 

34.3 

26.3 

1 1.1 

133.5 

64.2 

51.2 

(20) 

.28 

.54 

.057 

.09 

0 

.24 

.098 

.11 

.41 

.73 

.19 

.32 

.46 

1.09 

.69 

1.08 

32-37 

43-50 

3. 5-5.0 

2-3 

2 

2-6 

2. 0-3.0 

0-1 

33-39 

6-18 

33-36 

25-30 

7-16 

126-142 

59-68 

44-58 

B 

19.0 

21.8 

4.80 

3.2 

1.8 

2.0 

4.00 

0 

63.4 

16.2 

36.2 

24.8 

7.0 

235.0 

1 16.2 

58.0 

(5) 

.71 

.66 

.255 

.20 

.20 

0 

0 

0 

1.96 

.49 

.58 

.25 

.71 

3.10 

1.69 

3.03 

17-21 

20-24 

4.0-5. 5 

3-4 

1-2 

2 

4.0 

0 

60-71 

15-17 

35-38 

24-25 

6-9 

224-242 

1 12-122 

53-69 

C 

19.5 

21.5 

3.00 

3.0 

2.0 

2.5 

4.00 

0 

60.0 

15.5 

35.5 

22.0 

4.5 

216.5 

101.0 

53.5 

(2) 

1.50 

1.50 

0 

0 

0 

.50 

0 

0 

1.00 

.50 

.50 

0 

1.50 

3.50 

1.00 

4.50 

18-21 

20-23 

3.0 

3 

2 

2-3 

4.0 

0 

59-61 

15-16 

35-36 

22 

3-6 

213-220 

100-102 

49-58 

D 

31.3 

30.0 

5.58 

4.4 

2.3 

1.5 

4.80 

.5 

60.2 

16.0 

36.1 

26.4 

14.1 

241.5 

123.9 

67.9 

(25) 

.47 

.33 

.157 

.13 

.11 

.16 

.071 

.12 

.73 

.16 

.13 

.28 

.46 

1.15 

1.48 

1.64 

28-36 

27-35 

4. 0-7.0 

3-5 

1-3 

0-2 

4. 0-5.0 

0-2 

55-68 

15-17 

35-37 

24-29 

10-19 

231-251 

112-140 

52-85 

E 

25.8 

33.9 

4.50 

3.4 

2.0 

1.8 

2.20 

.7 

45.1 

23.1 

35.4 

25.5 

17.9 

181.8 

92.4 

55.1 

(33) 

.27 

.33 

.072 

.11 

.07 

.12 

.167 

.10 

.49 

1.13 

.23 

.17 

.41 

1.24 

1.06 

1.18 

23-30 

31-38 

3. 5-5.0 

2-5 

1-3 

0-3 

1 .0-4.0 

0-2 

41-54 

15-39 

33-37 

23-28 

13-23 

165-192 

78-104 

40-73 

F 

31.0 

35.0 

5.00 

4.0 

2.0 

3.0 

4.00 

0 

47.0 

17.0 

36.0 

28.0 

18.0 

205.0 

103.0 

53.0 

(D 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

31 

35 

5.0 

4 

2 

3 

4.0 

0 

47 

17 

36 

28 

18 

205 

103 

53 

G 

27.6 

29.1 

4.01 

3.0 

1.9 

2.0 

3.15 

.6 

48.3 

17.0 

35.9 

26.5 

14.5 

185.7 

95.2 

52.1 

(43) 

.37 

.36 

.107 

.08 

0 

.12 

.093 

.11 

.50 

.37 

.19 

.30 

.43 

1.95 

1.19 

.94 

24-35 

24-34 

3. 0-5.0 

2-4 

0-3 

0-4 

1.0-4. 5 

0-3 

43-56 

15-27 

34-39 

23-31 

9-23 

166-217 

83-113 

42-64 

H 

30.4 

17.9 

3.18 

2.9 

2.0 

2.2 

2.85 

.3 

45.1 

16.0 

35.9 

24.9 

7.7 

163.6 

85.5 

45.2 

(20) 

.49 

.45 

.083 

.06 

.10 

.08 

.064 

.11 

.46 

.18 

.15 

.33 

.45 

1.84 

1.00 

.89 

26-34 

15-22 

2. 5-4.0 

2-3 

1-3 

2-3 

2. 5-3. 5 

0-1 

42-49 

15-17 

35-38 

22-29 

4-12 

150-178 

77-94 

39-53 

Table  3.  Variation  among  30  scale  characters  for  nine  species  of  Lepidophyma  from  Mexico.  Presentation  as  in  Table  1. 

Species 

FPT 

LTR 

DBPVR 

IWD1 

1WV1 

PTMP 

DBPVT 

GC1IL 

GUL 

PVTL 

VL 

4TL 

4TLD 

DOR 

DAPVR 

PVR 

gaigeae 

33.5 

46.5 

4.03 

2.2 

2.0 

3.8 

2.54 

.5 

36.4 

11.3 

34.3 

26.3 

11.1 

133.5 

64.2 

51.2 

(20) 

.28 

.54 

.057 

.09 

0 

.24 

.098 

.11 

.41 

.73 

.19 

.32 

.46 

1.09 

.69 

1.08 

32-37 

43-50 

3. 5-4.0 

2-3 

2 

2-6 

2. 0-3.0 

0-1 

33-39 

6-18 

33-36 

25-30 

7-16 

126-142 

59-68 

44-58 

occulor 

19.2 

22.2 

4.44 

3.3 

1.9 

2.1 

4.1 1 

0 

63.2 

16.2 

36.1 

23.9 

6.8 

229.7 

1 1 1.9 

56.7 

(9) 

.46 

.55 

.306 

.17 

.11 

.11 

.11 1 

0 

1.28 

.36 

.35 

.44 

.70 

4.09 

3.04 

2.46 

17-21 

20-24 

3. 0-5. 5 

3-4 

1-2 

2-3 

4. 0-5.0 

0 

59-71 

15-18 

35-38 

22-25 

3-9 

213-242 

100-122 

49-69 

micropholis 

31.3 

30.0 

5.58 

4.4 

2.3 

1.5 

4.80 

.5 

60.2 

16.0 

36.1 

26.4 

14.1 

241.5 

123.9 

67.9 

(25) 

.47 

.33 

.157 

.13 

.11 

.16 

.071 

.12 

.73 

.16 

.13 

.28 

.46 

1.15 

1.48 

1.64 

28-36 

27-35 

4. 0-7.0 

3-5 

1-3 

0-2 

4. 0-5.0 

0-2 

55-68 

15-17 

35-37 

24-29 

10-19 

231-251 

112-140 

52-85 

sylvaticum 

27.6 

28.5 

4.01 

3.2 

2.0 

2.0 

2.78 

.6 

46.6 

18.9 

35.7 

25.8 

14.2 

180.0 

92.3 

51.7 

(98) 

.27 

.64 

.074 

.06 

.05 

.07 

.083 

.06 

.33 

.52 

.12 

.18 

.46 

1.36 

.77 

.70 

23-35 

15-38 

2.5-5. 0 

2-5 

0-3 

0-4 

1. 0-4.5 

0-3 

41-56 

15-39 

33-39 

20-31 

4-23 

1 50-2 1 7 

77-113 

39-73 

tuxtlae 

24.6 

34.1 

4.09 

3.7 

2.3 

5.5 

.29 

1.6 

42.6 

39.3 

39.6 

25.8 

16.3 

172.0 

79.6 

45.0 

(81) 

.22 

.20 

.053 

.06 

.06 

.11 

.032 

.09 

.28 

.43 

.14 

.16 

.22 

.97 

.70 

.43 

20-29 

30-40 

3. 0-5.0 

3-5 

0-3 

4-9 

0-.9 

0-4 

37-49 

30-47 

37-42 

23-28 

13-20 

150-190 

69-98 

37-55 

pajapanensis 

33.3 

40.5 

4.21 

3.4 

1.8 

7.5 

.14 

.1 

38.9 

43.8 

36.6 

28.3 

13.8 

164.3 

69.6 

44.0 

(19) 

.35 

.40 

.088 

.12 

.10 

.31 

.037 

.07 

.53 

1.02 

.17 

.24 

.50 

1.59 

1.22 

.84 

30-36 

37-43 

4. 0-5.0 

3-4 

1-2 

6-10 

0-.5 

0-1 

35-43 

39-51 

35-38 

26-30 

10-17 

152-175 

59-82 

37-49 

flavimaculatum 

33.7 

28.5 

4.45 

3.8 

2.1 

6.8 

2.99 

.9 

47.3 

18.6 

36.6 

27.3 

14.7 

197.7 

90.1 

54.3 

(49) 

.56 

.26 

.084 

.08 

.05 

.20 

.119 

.09 

.41 

.41 

.13 

.25 

.34 

1.64 

1.17 

1.00 

27-41 

25-32 

3.0-5. 5 

3-5 

1-3 

4-1  1 

.5-4.0 

0-2 

40-55 

15-27 

34-38 

23-31 

10-22 

173-221 

75-118 

43-69 

smithii 

23.1 

17.5 

4.21 

3.0 

2.1 

2.7 

3.18 

.7 

52.4 

16.8 

35.8 

25.2 

5.3 

194.0 

94.2 

48.7 

(36) 

.56 

.16 

.102 

.05 

.08 

.14 

.067 

.13 

.39 

.36 

.18 

.24 

.33 

1.43 

1.15 

.73 

18-29 

15-19 

3.0-5. 5 

2-4 

1-3 

2-4 

2. 0-4.0 

0-3 

46-57 

14-25 

33-38 

22-28 

2-9 

178-214 

81-1 10 

37-61 

tarascae 

17.0 

16.7 

2.67 

2.7 

2.0 

4.0 

2.50 

.7 

42.3 

16.0 

34.3 

22.7 

1.7 

146.7 

75.0 

44.3 

(3) 

.58 

.33 

.333 

.33 

0 

0 

0 

.33 

.67 

0 

.33 

.33 

.33 

1.67 

1.00 

1.20 

16-18 

16-17 

2. 0-3.0 

2-3 

2 

4 

2.5 

0-1 

41-43 

16 

34-35 

22-23 

1-2 

145-150 

74-77 

42-46 

8 Contributions  in  Science,  Number  349 


Bezy:  Systematics  of  Lepidophyma  in  Mexico 


Table  2.  Continued. 


Group 

PVS 

PVT1 

PVT2 

PVT3 

RPOL 

RPAW 

KPFML 

RPFL1 

RMW 

RNL 

RML 

RAPPSL 

RPNH 

RSLH 

A 

36.5 

14.6 

3.9 

0 

.208 

.407 

.491 

.405 

.094 

.199 

.127 

.052 

.744 

.745 

(20) 

1.97 

1.05 

.74 

0 

.0056 

.0168 

.0264 

.0367 

.0400 

.0076 

.0587 

.0323 

.0087 

.0155 

22-50 

8-22 

0-1 1 

0 

.15-.  26 

.27-54 

.28-.  71 

0-.63 

0-.62 

. 1 5-.  2 6 

0-.84 

0-.59 

.68-80 

.59-86 

B 

28.8 

29.2 

30.8 

20.2 

.169 

.555 

.716 

.716 

0 

.148 

0 

.022 

.736 

.873 

(5) 

3.34 

1.59 

.58 

.97 

.0151 

.0281 

.0233 

.0233 

0 

.0104 

0 

.0222 

.0314 

.0381 

21-39 

23-32 

30-33 

18-23 

. 13-.22 

.50-.66 

.67-.78 

.67-78 

0 

.1 1-.18 

0 

0-.11 

.61 — .78 

.76-96 

C 

22.0 

31.5 

30.0 

16.0 

.145 

.629 

.709 

.709 

0 

.090 

0 

0 

.776 

.805 

(2) 

4.00 

.50 

0 

0 

.0251 

.0838 

.0443 

.0443 

0 

.0059 

0 

0 

.0281 

.0216 

18-26 

31-32 

30 

16 

.12-.  17 

,57-,69 

.67-75 

.67-75 

0 

.08-.  10 

0 

0 

.75-81 

.78-.83 

D 

36.2 

31.7 

26.0 

16.2 

.163 

.585 

.557 

.563 

0 

.147 

0 

0 

.747 

1.146 

(25) 

1.99 

.72 

.96 

.32 

.0060 

.0592 

.0175 

.0187 

0 

.0067 

0 

0 

.0133 

.0334 

17-56 

25-40 

17-35 

13-20 

. 1 2-.24 

,27-,85 

.42-.76 

.42-76 

0 

. 10-.22 

0 

0 

.66-. 95 

.94-1.76 

E 

24.3 

30.8 

22.6 

2.5 

.183 

.519 

.527 

.462 

.127 

.167 

.100 

.054 

.838 

1.170 

(33) 

1.50 

.70 

.84 

.53 

.0100 

.0283 

.0193 

.0214 

.0432 

.0088 

.0376 

.0356 

.0202 

.0252 

7-42 

22-37 

16-33 

0-13 

.07-.32 

.30-.75 

.27-.82 

. 19-.8 1 

0-.94 

.09-.28 

0-.85 

0-1.12 

.71-1.37 

.84-1.55 

F 

20.0 

33.0 

22.0 

9 

.200 

.466 

1.071 

.283 

.729 

.106 

.739 

0 

.750 

1.141 

< i ) 

0 

0 

0 

0 

0 

.0193 

0 

0 

0 

0 

0 

0 

0 

0 

20 

33 

22 

9.0 

.20 

.46 

1.07 

.28 

.73 

.11 

.74 

0 

.75 

1.14 

G 

23.7 

28.4 

2.09 

12.6 

.180 

.540 

.560 

.368 

.237 

.165 

.338 

.072 

.772 

1.171 

(43) 

1.30 

.65 

.73 

.50 

.0062 

0 

.0190 

.0309 

.0508 

.0073 

.0746 

.0473 

.0185 

.0181 

12-43 

19-35 

15-34 

4-18 

.1 1-.27 

.4Q-.69 

.36-.  81 

0-.71 

0-.84 

.07-.29 

0-1.32 

0-1.65 

.64-1.40 

.86-1.41 

H 

14.2 

30.3 

21.3 

15.9 

.220 

.633 

.464 

.406 

.088 

.156 

.074 

.044 

.749 

1.073 

(20) 

.78 

.50 

.87 

.18 

.0194 

.0120 

.0311 

.0370 

.0399 

.0121 

.0373 

.0443 

.0368 

.0377 

7-20 

26-34 

15-28 

15-17 

. 1 4-56 

.39-.82 

.1 1-.70 

.1  1-.65 

0-.56 

.09-28 

0-.63 

0-.89 

.65-1.39 

.89-1.61 

Table  3.  Continued. 

Species 

PVS 

PVT  1 

PVT2 

PVT3 

RPOL 

RPAW 

RPFML 

RPFL1 

RMW 

RNL 

RML 

RAPPSL 

RPNH 

RSLH 

gaigeae 

36.5 

14.6 

3.9 

0 

.208 

.407 

.491 

.405 

.094 

.199 

.127 

.052 

.744 

.745 

(20) 

1.97 

1.05 

.74 

0 

.0056 

.0168 

.0264 

.0367 

.0400 

.0076 

.0587 

.0323 

.0087 

.0155 

22-50 

8-22 

0-1 1 

0 

. 1 5-.  26 

.27-.54 

.28—.  7 1 

0-.63 

0-1.49 

. 1 5-26 

0-.84 

0-.59 

.68-. 80 

.59-. 86 

occulor 

26.9 

29.9 

30.6 

19.0 

.166 

.576 

.714 

.714 

0 

.131 

0 

.016 

.748 

.854 

(9) 

2.76 

1.18 

.43 

1.02 

.0099 

.0271 

.0188 

.0188 

0 

.0129 

0 

.0159 

.0237 

.0295 

18-39 

23-32 

30-33 

16-23 

. 1 2-,22 

.50-.69 

.66-.  78 

.67-.77 

0 

oo 

OO 

o 

0 

0-.1 1 

.6 1 —.8 1 

.76-. 96 

micropholis 

36.2 

31.7 

26.0 

16.2 

.163 

.585 

.557 

.563 

0 

.147 

0 

0 

.747 

1.146 

(25) 

1.99 

.72 

.95 

.32 

.0060 

.0283 

.0175 

.0187 

0 

.0067 

0 

0 

.0133 

.0334 

17-56 

25-40 

17-35 

13-20 

.12-.24 

.27-85 

.42-.76 

,42-,76 

0 

.09-.22 

0 

0 

,66-.95 

.94-1.76 

sylvaticum 

21.9 

29.7 

21.0 

9.8 

.190 

.551 

.532 

.408 

.171 

.165 

.200 

.059 

.790 

1.152 

(98) 

.88 

.40 

.47 

.62 

.0062 

.0109 

.0141 

.0172 

.0287 

.0052 

.0373 

.0253 

.0133 

.0144 

7-43 

19-37 

15-34 

0-18 

.07-.  56 

.30-. 82 

.1  1-1.07 

0 — .8 1 

0-.95 

.07-.31 

0-1.32 

0-1.65 

.64-1.40 

.84-1.61 

tuxtlae 

5.1 

39.9 

32.2 

1.7 

.224 

.830 

.734 

.042 

1.024 

.108 

1.216 

.027 

.734 

.547 

(81) 

.47 

.37 

.63 

.33 

.0047 

.0128 

.0092 

.0126 

.0243 

.0049 

.0539 

.0159 

.0054 

.0106 

0-18 

31-49 

1 1-42 

0-15 

.1  1-.33 

.59-1.12 

O' 

OO 

f 

OO 

T 

0-.57 

0-1.31 

.04-.29 

0-3.15 

0-.99 

.63-.83 

.32-77 

pajapanensis 

2.8 

41.2 

26.6 

.1 

.222 

.540 

.803 

.157 

1.087 

.163 

1.656 

.002 

.764 

.485 

(19) 

.56 

1.02 

1.33 

.06 

.0081 

.0189 

.0131 

.0246 

.0264 

.0113 

.1192 

.0015 

.0116 

.0349 

0-8 

34-49 

15-40 

0-1 

. 1 3-28 

.38-.69 

.70-.90 

0-.31 

.85-1.29 

. 1 0— .28 

1.00-2.65 

0-.03 

.69-86 

,32-.88 

Jlavimaculatum 

20.1 

34.2 

25.3 

13.7 

.204 

.411 

.699 

.175 

.834 

.145 

.978 

.061 

.736 

.557 

(49) 

1.17 

.65 

.71 

.76 

.0089 

.0199 

.0126 

.0128 

.0447 

.0075 

.0687 

.0356 

.0063 

.0154 

6-44 

24-43 

17-35 

1-24 

.11-39 

.19-69 

,52-,88 

0-.73 

0-1.31 

.07-.28 

0-2.43 

0-1.66 

.58-83 

.36-89 

smithii 

17.0 

31.1 

26.9 

16.3 

.181 

.723 

.713 

.349 

.608 

.137 

.783 

.099 

.794 

.904 

(36) 

.75 

.45 

.60 

.4! 

.0065 

.0459 

.0141 

.0414 

.0681 

.0052 

.1050 

.0398 

.0093 

.0106 

8-29 

25-39 

18-32 

13-27 

.1  1-.28 

.43-1.35 

.54— .89 

0-.79 

0-1.07 

.08-24 

0-2.62 

0-1.04 

.66-.92 

.61-1.22 

tarascae 

13.7 

30.7 

25.0 

16.0 

.299 

.591 

.361 

.361 

0 

.154 

0 

0 

.982 

.756 

(3) 

1.33 

.33 

1.53 

0 

.0717 

.0923 

.0664 

.0664 

0 

.0130 

0 

0 

.0227 

.0417 

1 1-15 

30-31 

23-28 

16 

.19-.44 

.41-.71 

.29-49 

.29-49 

0 

.14-18 

0 

0 

.94-1.02 

.69-. 83 

Contributions  in  Science,  Number  349 


Bezy:  Systematics  of  Lepidophyma  in  Mexico  9 


in  PTMP  (0-3  in  95/97  EFGH  and  4-11  in  all  J)  and  RSLH 
(0.84-1 .60  in  all  EFGH  and  0.36-0.79  in  49/50  J)  (Table  2). 
The  two  specimens  of  EFGH  with  a PTMP  of  4 are  from  a 
population  (6)  at  the  northern  end  of  the  range  (Fig.  1),  and 
the  specimen  of  J with  RSLH  of  0.89  is  from  a population 
in  Tabasco  (37),  that  is  separated  from  the  southernmost 
sample  (12)  of  EFGH  by  350  km  and  by  intervening  pop- 
ulations that  are  clearly  assigned  to  J.  Thus  the  populations 
of  EFGH  and  J that  are  most  similar  in  morphology  are  not 
geographically  intermediate,  and  the  multivariate  and  uni- 
variate ditferences  between  the  two  population  groups  are 
sufficiently  constant  that  they  are  judged  to  represent  units 
that  are  likely  reproductively  isolated. 

Group  D is  multivariately  closest  to  G (Fig.  2).  The  four 
populations  of  D are  from  the  Sierra  del  Abra  ofTamaulipas 
and  San  Luis  Potosi  (Fig.  6),  and  differ  in  DOR  from  all 
populations  of  group  EFGH  including  those  in  the  Sierra 
Madre  Oriental  to  the  west  and  Sierra  Tamaulipas  to  the  east 
(Table  2).  While  two  of  the  three  specimens  from  the  Sierra 
Tamaulipas  (sample  7,  group  G)  approach  group  D in  dis- 
criminant space,  they  are  not  geographically  intermediate 
(Fig.  6)  and  do  not  bridge  the  gap  between  the  two  in  DOR 
(Table  2).  It  is  concluded  that  group  D should  be  considered 
specifically  distinct  from  EFGH. 

Groups  B and  C are  closest  to  D in  discriminant  space, 
but  differ  in  FPT  (18-21  vs.  28-36)  and  LTR  (20-24  vs.  27- 
35).  Evaluation  of  these  differences  is  hampered  by  small 
sample  sizes  of  B (N  = 4)  and  C (N  = 2).  As  an  estimate  of 
the  range  of  variation  that  would  be  expected  with  larger 
samples,  three  standard  deviations  of  D were  added  to  and 
subtracted  from  the  means  of  B and  C.  The  estimated  ranges 
of  B and  C overlap  each  other  for  all  characters  but  differ 
from  the  observed  range  ofD  for  FPT  (12-27  vs.  28-36)  and 
LTR  (17-26  vs.  27-35).  The  populations  of  B and  C are 
located  in  the  Jalpan  Valley  of  Queretaro  and  San  Luis  Potosi 
(Fig.  6),  and  their  combined  ranges  of  variation  differ  in  six 
characters  from  the  populations  of  E occurring  in  the  Sierra 
Madre  Oriental,  22  km  to  the  east.  While  larger  sample  sizes 
are  necessary  to  fully  evaluate  the  differences  between  groups 
B and  C,  their  multivariate  juxtaposition  and  the  small  uni- 
variate differences  between  them  indicate  they  are  probably 
members  of  the  same  species.  On  the  other  hand  the  number 
and  magnitude  of  the  univariate  differences  between  BC  and 
its  nearest  geographic  (E)  and  discriminant  (D)  neighbors  are 
such  that  they  are  not  likely  to  be  bridged  by  larger  samples. 
Group  BC  is  considered  specifically  distinct  from  D and 
EFGH. 

Group  A is  strongly  separated  in  discriminant  space  from 
the  populations  of  all  other  groups  (Figs.  2-3).  It  is  morpho- 
logically and  geographically  closest  to  group  E (Fig.  6),  but 
differs  in  6 of  the  30  characters  (Table  2).  The  univariate  and 
multivariate  differences  between  A and  other  population 
groups  clearly  qualify  it  as  a distinct  morphospecies. 

While  the  combined  problems  of  small  sample  sizes,  dis- 
junct distributions,  and  discordant  variation  confound  some 
of  the  decisions,  four  unique  morphological  units  of  Lepi- 
dophyma  are  recognizable  in  northeastern  Mexico:  A,  BC, 
D,  and  EFGH. 


ALLOCATION  OF  NAMES 

Several  of  the  groups  identified  in  the  discriminant  analyses 
include  lizards  that  are  either  types  or  are  from  or  near  the 
type  locality  of  named  taxa:  Group  A:  Population  21:  L. 
gaigeae  Mosauer,  1936;  B:  20:  L.  smithu  occulor  Smith,  1942; 
D:  14:  L.  micropholis  Walker,  1955;  E:  11:  L.  sylvaticum 
Taylor,  1939;  G:  6:  L.  flavimaculatum  tenebrarum  Walker, 
1955;  I:  51:  L.  tuxtlae  Werler  and  Shannon,  1957,  41:  L. 
pajapanensis  Werler,  1957,  48:  L.  sawini  Smith,  1973,  47: 
L.  alvarezi  Smith,  1 973;  J:  28:  L.  flavimaculatum  A.  Dumeril 
in  Dumeril  and  Dumeril,  1851;  K:  44:  L.  smithii  Bocourt, 
1876;  L:  46:  L.  tarascae  Bezy,  Webb,  and  Alvarez,  1982. 

The  oldest  available  names  for  the  species  units  recognized 
in  northern  Mexico  (Fig.  9)  from  the  foregoing  discussions 
are:  A,  L.  gaigeae ; BC,  L.  occulor,  D,  L.  micropholis-,  and 
EFGH,  L.  sylvaticum.  These  are  summarized  below. 

The  systematic  relationships  among  populations  of  Lepi- 
dophyma  in  southern  Mexico  currently  are  under  study  (Bezy, 
in  prep.);  the  southern  population  groups  used  in  this  paper 
are  considered  to  represent  the  following  species:  I — L.  pa- 
japanensis (Veracruz)  and  L.  tuxtlae  (Veracruz,  Oaxaca, 
Chiapas);  J = L.  flavimaculatum  (Atlantic  versant  east  of  the 
Isthmus  ofTehuantepec  in  Oaxaca,  Veracruz,  Tabasco,  Chia- 
pas, Quintana  Roo);  K = L.  smithii  (Pacific  versant  of  Guer- 
rero, Oaxaca,  Chiapas);  and  L = L.  tarascae  (coastal  Mi- 
choacan). 

SPECIES  ACCOUNTS 
Lepidophyma  gaigeae  Mosauer 
Group  A;  Figures  7,  10 

Lepidophyma  gaigeae  Mosauer,  1936:3.  Holotype:  MCZ 

42145:  Durango,  State  of  Hidalgo,  Mexico. 

Gaigeia  gaigeae:  Smith,  1939:24. 

DIAGNOSTIC  CHARACTERS.  Differs  from  other 
members  of  the  genus  except  L.  radula  in  having  43-50 
subequal  scales  (rather  than  1 5-42  discrete  rows  of  enlarged 
tubercles)  along  the  side  of  the  body  (axilla  to  groin)  (Figs. 
7-8)  and  fewer  dorsal  scales  (126-142  vs.  145-251)  (Table 
3).  It  differs  from  L.  radula  and  L.  dontomasi  in  having  two 
(rather  than  one)  caudal  interwhorls  complete  ventrally. 

DISTRIBUTION.  The  species  is  known  from  Hidalgo  (near 
the  type  locality)  and  Queretaro  (between  El  Lobo  and  Jalpan; 
Dixon  et  al.,  1972),  where  it  occurs  in  limestone  crevices 
primarily  in  pine-oak  woodland  (Fig.  9). 

REMARKS.  The  high  degree  of  separation  of  L.  gaigeae 
from  other  populations  in  the  discriminant  analyses  is  con- 
sistent with  its  proposed  separate  generic  (Smith,  1942)  or 
subgeneric  (Smith,  1973)  status.  Geographic  variation  and 
relationships  of  this  form  to  L.  dontomasi  and  L.  radula  are 
currently  under  study  (Bezy,  in  prep.). 

KARYOTYPE.  Lepidophyma  gaigeae  has  a diploid  chro- 
mosome number  of  38  with  nine  pairs  of  macrochromo- 
somes and  10  pairs  of  microchromosomes.  The  karyotype  is 
unique  in  the  genus,  but  closest  to  those  of  L.  flavimaculatum, 
L.  pajapanensis.  and  L.  tuxtlae  (Bezy,  1972). 


10  Contributions  in  Science,  Number  349 


Bezy:  Systematics  of  Lepidophyma  in  Mexico 


Figure  9.  Distribution  of  the  four  species  of  Lepidophyma  recog- 
nized in  northeastern  Mexico  (north  of  19°N).  Lines  enclose  the 
samples  included  in  each  species. 


Lepidophyma  occulor  Smith 

Group  BC;  Figure  10 

Lepidophyma  smithii  occulor  Smith,  1942:378.  Holotype: 

USNM  47133:  Jalpan,  Queretaro. 

Lepidophyma  flavimaculatum  occulor:  Walker,  1955:5. 
Lepidophyma  occulor:  Bezy,  1972:15. 

DIAGNOSTIC  CHARACTERS.  Differs  from  all  other 
species  of  Lepidophyma  except  L.  micropholis  in  having  more 
gulars  (59-71  vs.  33-57),  and  from  L.  micropholis  in  having 
fewer  femoral  pores  (17-21  vs.  28-36),  fewer  lateral  tubercle 
rows  (20-24  vs.  27-35),  and  fewer  divided  fourth  toe  lamellae 
(3-9  vs.  10-19)  (Table  3). 

DISTRIBUTION.  Lepidophyma  occulor  is  known  from 
four  localities  in  the  Jalpan  Valley  of  Queretaro  and  San  Luis 
Potosi  (Fig.  9),  where  it  has  been  found  beneath  stones  in 
arid  tropical  scrub  (Dixon  et  al.,  1972). 

REMARKS.  The  species  alternatively  has  been  considered 
a subspecies  of  either  L.  smithii  (Smith,  1942)  or  L.  Jlavi- 
maculatum  (Walker,  1955)  from  both  of  which  it  is  well 
separated  multivariately  (Fig.  3),  differing  from  the  former 
in  lateral  tubercle  rows  and  gulars  and  from  the  latter  in 
femoral  pores,  lateral  tubercle  rows,  pretympanics,  gulars, 
and  divided  fourth  toe  lamellae.  It  is  multivariately  closest 
to  L.  micropholis  from  which  it  differs  in  femoral  pores. 


Figure  10.  Living  individuals  of  Lepidophyma  gaigeac  (upper, 
LACM  127170),  L.  occulor  (middle,  sample  18,  TCWC  35605),  and 
L micropholis  (lower,  sample  16.  TCWC  60767). 


lateral  tubercle  rows,  and  divided  fourth  toe  lamellae  (Table 
3). 

KARYOTYPE.  Lepidophyma  occulor  has  a diploid  chro- 
mosome number  of  36,  with  the  lowest  number  of  micro- 
chromosomes (18)  known  in  the  family  Xantusiidae  (Bezy, 
1972). 

Lepidophyma  micropholis  Walker 

Group  D;  Figure  10 

Lepidophyma  micropholis  Walker,  1955:6.  Holotype:  UMMZ 
101298:  cave  at  El  Pachon,  about  5 miles  NNE  of  Antigua 
Morelos,  Tamaulipas. 

DIAGNOSTIC  CHARACTERS.  Differs  from  all  other 
species  in  the  genus  (except  L.  occulor)  in  having  more  dorsal 


Contributions  in  Science,  Number  349 


Bezy:  Systematics  of  Lepidophyma  in  Mexico  1 


! 


Figure  1 1.  Living  individuals  of  southern  Madrean  (upper,  sample 
1 1 . LACM  106752)  and  northern  Madrean  (lower,  sample  8,  LACM 
131 146)  population  groups  of  Lepidophyma  sylvalicum. 


scales  (231-251  vs.  126-222)  and  from  L.  occulor  in  having 
more  lateral  tubercle  rows  (27-35  vs.  20-24)  (Table  3). 

DISTRIBUTION.  Lepidophyma  micropholis  occurs  in 
southern  Tamaulipas  and  northern  San  Luis  Potosi  at  four 
localities  situated  along  the  Sierra  del  Abra  (Fig.  9).  This 
cavemiferous,  low-lying  range  constitutes  the  easternmost 
front  of  the  Sierra  Madre  Oriental  in  the  highly  dissected 
region  between  the  Rio  Guayalejo  and  the  Rio  Tamuin 
(Mitchell  et  al.,  1 977).  The  lizards  have  been  found  primarily 
in  limestone  caves  (El  Pachon  and  Quintero)  and  fissures. 

REMARKS.  This  extensively  cavernicolous  species  is 
closest  in  scalation  to  L.  occulor  and  L.  sylvaticum,  and  future 
work  may  demonstrate  the  existence  of  morphologically  and 
biochemically  intermediate  populations,  similar  to  those  oc- 
curring between  the  epigean  and  troglodytic  Astyanax  mex- 
icanus  of  the  Sierra  del  Abra  (Avise  and  Selander,  1972; 
Mitchell  et  al.,  1977). 

KARYOTYPE.  Lepidophyma  micropholis  has  a diploid 
chromosome  number  of  36  with  1 6 macrochromosomes  and 
20  microchromosomes  (Bezy,  1972). 

Lepidophyma  sylvaticum  Taylor 

Group  EFGH;  Figures  7-8,  11-12 

Lepidophyma  sylvatica  Taylor,  1 939: 131.  Holotype;  FMNH 

100102:  7 mi.  north  of  Zacaultipan,  Hidalgo. 

Gaigeia  sylvatica:  Smith,  1942:380. 


Figure  12.  Living  individuals  of  northern  Madrean  (upper,  sample 
6.  LACM  106752)  and  western  (lower,  sample  1,  LACM  106781) 
population  groups  of  Lepidophyma  sylvaticum. 


Lepidophyma  sylvaticum:  Walker,  1955:9. 

Lepidophyma  flavimaculatum  tenebrarum  Walker,  1955:1. 
NEW  SYNONYMY.  Holotype:  UMMZ  101374:  ±5  miles 
NW  (by  road)  of  Gomez  Farias  in  the  Sierra  Madre  Ori- 
ental at  “Rancho  del  Cielo.” 

DIAGNOSTIC  CHARACTERS.  Differs  from  L.  gaigeae 
and  L.  occulor  in  numbers  of  gulars  (41-56  vs.  33-39  and 
59-7 1 , respectively),  from  L.  micropholis  in  numbers  of  dor- 
sals (150-217  vs.  231-251),  from  L.  tarascae  in  numbers  of 
femoral  pores  (23-35  vs.  16-18),  from  L.  smithii  in  having 
a parietal  foramen,  from  L.  flavimaculatum  in  numbers  of 
pretympanics  (0-3  vs.  4-11,  99%)  and  ratio  of  supralabial 
height  (0.84-1.61  vs.  0.36-0.79,  99%),  from  L.  tuxtlae  in 
ratio  of  supralabial  height  (0.84-1.61  vs.  0.32-0.77),  and 
from  L.  pajapanensis  in  numbers  of  pretympanics  (0-4  vs. 
6-10)  (Table  3). 

DISTRIBUTION.  The  12  populations  occur  from  Vera- 
cruz to  Nuevo  Leon  along  the  Sierra  Madre  Oriental  and 
adjacent  Mesa  Central  and  Sierra  Tamaulipas  (Fig.  9). 

REMARKS.  The  four  population  groups  included  in  L. 
sylvaticum  are  moderately  divergent  from  one  another  and 
further  work  may  indicate  that  one  or  more  of  them  should 
be  given  separate  taxonomic  recognition  (i.e.,  subspecies). 

The  northern  Madrean  group  (G)  includes  four  populations 
in  southern  Tamaulipas  and  northern  San  Luis  Potosi,  three 
from  along  the  main  axis  of  the  Sierra  Madre  Oriental  and 


12  Contributions  in  Science,  Number  349 


Bezy:  Systematics  of  Lepidophyma  in  Mexico 


AX 

6 + 8 


AK  NA 

4 5 

a*  • • V • 


KK 

7 9 


XX  /#/S  DM  Xa  *K  /lt*» 


Figure  13.  Karyotypes  of  Lepidophyma  sylvaticum  (sample  6;  upper,  LACM  106758,  <5;  lower,  LACM  106763,  9). 


one  in  the  Sierra  Tamaulipas.  They  differ  from  the  three 
populations  (group  E)  of  southern  San  Luis  Potosi,  Quere- 
taro,  and  Hidalgo  in  lateral  tubercle  rows  (34/39  northern 
Madrean  with  31  or  less,  30/33  southern  Madrean  with  32 
or  more)  (Table  2).  The  hiatus  between  the  ranges  of  the  two 
corresponds  roughly  to  the  Rio  Panuco  gap  in  the  Sierra 
Madre  Oriental.  Should  further  work  demonstrate  a need  to 
accord  them  separate  nomenclatural  status,  L.  sylvaticum 
Taylor,  1939  is  applicable  to  the  southern  group,  and  L.  f 
tenebrarum  Walker,  1 955  is  available  for  the  northern  group. 

The  southernmost  specimen  (group  F)  assigned  to  L.  syl- 
vaticum is  from  ca.  170  km  SE  of  the  type  locality  and  occurs 
at  the  northern  base  of  the  Cordillera  Volcanica  in  central 
Veracruz  (Fig.  6).  It  has  similarities  to  both  southern  and 
northern  Madrean  L.  sylvaticum,  being  closest  to  the  former 
in  lateral  tubercle  rows  and  femoral  pores,  and  to  the  latter 
in  dorsals.  No  other  Lepidophyma  are  known  from  the  Cor- 
dillera Volcanica.  The  nearest  populations  to  the  south  are 
L.  tuxtlae  and  L.  pajapanensis  of  the  Tuxtlas  region  (which 
differ  from  L.  sylvaticum  in  numbers  of  large  paravertebrals, 
pretympanics,  and  dorsals,  and  in  ratio  of  supralabial  height), 
and  L.  flavimaculatum  of  the  northern  Isthmus  of  Tehuan- 
tepec (which  differ  in  pretympanics,  lateral  tubercle  rows, 
and  ratio  of  supralabial  height)  (Tables  2-3). 

The  western  group  of  L.  sylvaticum  (H,  Figs.  6,  8,  12) 
includes  one  population  in  the  Sierra  Alvarez  on  the  Mesa 
Central  of  southeastern  San  Luis  Potosi  and  two  in  canyons 
around  the  northern  base  of  the  Sierra  Madre  Oriental  below 
the  Cumbres  de  Monterey  of  Nuevo  Leon.  A fourth  locality 
(sample  3)  is  represented  by  a fragmentary  specimen,  but  the 
limited  data  obtainable  from  it  suggests  it  is  a member  of 
the  western  group.  The  group  differs  from  all  other  L.  syl- 
vaticum in  lateral  tubercle  rows,  and  from  southern  Madrean 


samples  of  L.  sylvaticum  in  numbers  of  fourth  toe  lamellae 
and  numbers  of  paravertebrals  larger  than  three  dorsal  scales 
(Table  2). 

KARYOTYPE.  Chromosomal  information  was  obtained 
from  16  specimens  (66,  92,  1 juv.)  of  L.  sylvaticum'.  three 
from  sample  1 1,  one  from  10,  eight  from  6,  and  four  from 
1.  A total  of  271  metaphase  spreads  were  studied. 

In  all  four  populations  the  karyotype  was  found  to  consist 
of  a diploid  number  of  36  with  16  macrochromosomes  and 
20  microchromosomes  (Fig.  13).  There  are  five  metacentric 
to  submetacentric  (Nos.  I,  2,  2A,  5,  7),  two  subtelocentric 
(3,  4),  and  one  acrocentric  (9)  pairs  of  macrochromosomes 
(pair  numbering  after  Bezy,  1972).  No  secondary  constric- 
tions were  observed.  The  cells  of  six  specimens  (52,  1 juv.) 
of  sample  6 were  found  consistently  to  have  a pair  of  het- 
eromorphic  chromosomes  involving  the  largest  pair  of  mi- 
crochromosomes, with  a metacentric  member,  ca.  1.5  times 
the  size  of  the  next  largest  micro  (Fig.  1 3). 

The  karyotype  of  L.  sylvaticum  appears  identical  in  all 
respects  (except  the  heteromorphism)  to  that  of  L.  micro- 
pholis  (Bezy,  1972).  It  differs  from  that  of  L.  flavimaculatum, 
L.  tuxtlae,  and  L.  pajapanensis  in  having  one  less  pair  of 
macrochromosomes  (the  large  metacentric  2A  presumably 
was  formed  from  centric  fusions  involving  pairs  6 and  7),  a 
pair  3 that  lacks  terminal  satellites,  and  a submetacentric 
rather  than  subtelocentric  pair  7;  from  that  of  L.  smithii  in 
that  pair  2A  is  more  metacentric  (rather  than  submetacen- 
tric), pair  3 lacks  satellites,  and  the  smallest  macro  pair  is 
acrocentric  rather  than  subtelocentric;  and  from  that  of  L. 
occulor  in  having  one  more  pair  of  macrochromosomes,  one 
less  pair  of  microchromosomes,  and  an  acrocentric  (rather 
than  submetacentric)  pair  9. 

Lepidophyma  sylvaticum  thus  differs  karyotypically  from 


Contributions  in  Science,  Number  349 


Bezy:  Systematics  of  Lepidophyma  in  Mexico  13 


the  species  that  are  closest  to  it  in  scalation  (L.  flavimacu- 
latum,  L.  smithii,  and  L.  occulor)  except  L.  micropholis.  Such 
chromosomal  differences  in  themselves  would  not  be  ex- 
pected to  present  reproductive  barriers,  and  some  cases  of 
extraordinary  geographic  variation  in  karyotypes  have  been 
documented  in  species  of  other  lizard  families  (e.g..  Hall  and 
Selander,  1973;  Sites,  1983).  However,  among  lizards  chro- 
mosomal divergence  most  often  is  associated  with  differen- 
tiation at  or  above  the  species  level.  The  karyotypic  identity 
of  L.  micropholis  and  L.  sylvaticum  serves  to  underscore  the 
morphologic  and  biogeographic  relationships  which  suggest 
that  the  former  may  be  a troglodytic  derivative  of  the  latter, 
and  that  future  work  might  demonstrate  a morphologic  and 
genetic  continuum  between  the  two. 

The  microchromosomal  heteromorphism  found  in  the  fe- 
males of  sample  6 was  not  detected  in  samples  1,  10,  and 
1 1.  It  occurs  in  all  females  (five;  plus  one  juvenile)  of  sample 
6,  but  is  absent  in  the  two  females  of  sample  1 and  in  all 
males  studied  (two  each  from  samples  1,  6,  and  1 1,  and  one 
from  1 0).  It  may  constitute  a sex  chromosomal  heteromorph- 
ism (ZW)  present  in  population  6,  absent  in  population  1, 
and  of  unknown  occurrence  in  populations  10  and  11  (no 
preparations  from  females  available).  On  the  other  hand,  it 
may  represent  a heterozygous  condition  where  unsampled 
homozygous  individuals  (for  the  large  macrochromosome) 
occur  in  the  population,  or  in  adjacent  populations.  Until 
additional  material  is  obtained,  all  that  can  be  said  is  that  it 
is  a heteromorphic  condition  which  has  been  found  only  in 
females  of  population  6,  and  not  detected  in  any  other  pop- 
ulation in  the  family. 

SEX  RATIO.  Two  samples  of  L.  sylvaticum  have  sex  ra- 
tios (2/<3  + 2)  that  differ  significantly  (0.05  level,  Fisher  exact 
test,  Yates  correction)  from  0.50:  sample  6 with  0.89  (25/ 
28)  and  sample  10  with  0.85  (1 1/13).  The  skewed  sex  ratios 
of  these  samples  have  been  discussed  earlier  in  relationship 
to  the  unisexual  populations  of  L.  flavimaculatum  occurring 
in  Panama  and  Costa  Rica  (Bezy,  1 972).  Although  the  sample 
sizes  of  populations  6 and  10  of  L.  sylvaticum  remain  smaller 
than  desirable,  both  are  now  sufficiently  large  to  conclude 
that  the  sex  ratios  differ  significantly  (0.05  level)  from  those 
of  other  Lepidophyma  populations  in  eastern  Mexico:  L. 
tuxtlae  (30/59)  and  L.  gaigeae  ( 1 50/260). 

In  addition  to  previously  discussed  factors  that  may  be 
responsible  for  the  observed  skewed  sex  ratios  (Bezy,  1972), 
temperature-dependent  sex  determination  has  now  been  doc- 
umented for  lizards  (Bull,  1980),  and  comparable  effects  could 
be  operative  in  Lepidophyma.  Theoretical  considerations 
would  predict  that  this  may  not  be  the  case  in  that  temper- 
ature-dependent sex  determination  is  thought  to  interfere 
with  the  evolution  of  both  viviparity  (ovoviviparity)  and  sex 
chromosomes  (Bull,  1980).  Viviparity  appears  to  be  univer- 
sal among  xantusiids  (Blackburn,  1982),  and  the  population 
(6)  of  L.  sylvaticum  with  the  most  aberrant  sex  ratio  is  the 
only  one  in  the  family  with  heteromorphism,  possibly  in- 
dicating the  presence  of  sex  chromosomes.  Experimental  data 
are  needed  to  evaluate  the  role  of  environmental  factors, 
particularly  temperature,  in  determining  sex  ratio  in  xantu- 
siids. 

The  aberrant  sex  ratios  and  the  heteromorphic  chromo- 

14  Contributions  in  Science,  Number  349 


somes  of  populations  of  L.  sylvaticum  could  be  a conse- 
quence of  hybridization.  Both  conditions  frequently  are  found 
in  hybrid  populations,  the  skewed  sex  ratios  perhaps  resulting 
from  a disruption  in  the  balance  of  the  sex-determining 
mechanism  or  an  increased  expression  of  lethals  in  one  of 
the  sexes  (Darevsky  et  al.,  1978;  White,  1973).  The  identi- 
fication of  populations  homozygous  for  the  large  microchro- 
mosome and  additional  data  (e.g.,  allozymes)  would  be  re- 
quired to  establish  the  existence  of  and  possible  participants 
in  such  a hybridization. 

KEY  TO  THE  SPECIES  OF  LEPIDOPHYMA 
OF  NORTHEASTERN  MEXICO 

la.  Side  of  body  lacking  vertical  rows  of  enlarged  keeled 
tubercles  (Fig.  7),  but  with  43-50  subequal  scales  (axilla 
to  groin);  less  than  145  dorsal  scales  (occiput  to  rump); 
tail  with  two  interwhorls  complete  dorsally  and  ventrally 

/..  gaigeae 

lb.  Side  of  body  with  enlarged,  keeled  tubercles  arranged  in 
1 5-42  vertical  rows  (A-G)  separated  by  smaller  granular 
scales  (Figs.  7-8);  145  or  more  dorsal  scales  (O-R);  tail 
usually  with  more  than  2 mterwhorls  complete  dorsally 

2 

2a.  Total  femoral  pores  2 1 or  less  (gulars  59  or  more;  divided 
4th  toe  lamellae  9 or  less;  lateral  tubercle  rows  24  or 


less) L.  occulor 

2b.  Total  femoral  pores  23  or  more  3 

3a.  Dorsal  scales  (O-R)  231  or  more  (gulars  55  or  more; 
divided  4th  toe  lamellae  1 0 or  more;  lateral  tubercle  rows 
27  or  more) L.  micropholis 


3b.  Dorsal  scales  (O-R)  2 1 7 or  less  (gulars  56  or  less;  divided 
4th  toe  lamellae  4-23;  lateral  tubercle  rows  15-38)  . . . 
L.  sylvaticum 

SPECIMENS  EXAMINED 

The  351  specimens  and  52  population  samples  studied  from 
Mexico  are  listed  below.  Sample  numbers  are  in  parentheses 
preceding  localities. 

L.  gaigeae 

HI DALGO:  (2 1 ):  La  Placita,  8 km  S Jacala  (UIMNH  26 1 80- 
86,  26191-99,  26204,  26207-09). 

L.  micropholis 

SAN  LUIS  POTOSI:  (15):  6 mi.  E Valles  (BCB  13837-42); 
(16):  5.5  mi.  S,  1.4  mi.  E Valles  (TCWC  60621,  60766-67). 
TAMAULIPAS:  (13):  Gruta  de  Quintero,  1.5  mi.  S Quintero 
(AMNH  93409,  LACM  66662,  SAM  885);  (14):  cave  at  El 
Pachon,  ca.  5 mi.  (by  rd)  NNE  Antigua  Morelos  (LACM 
106767-68.  UAZ  28762,  28767-69,  UMMZ  101299, 
102886-88);  11.3  mi.  S Ciudad  Mante,  Hwy  85  (TCWC 
57256). 

L.  occulor 

QUERETARO:  (18):  2.5  mi.  S Conca,  Hda.  Conca  (TCWC 
35605-06,  48499);  (19):  Jalpan  (LJSNM  47134-35);  (20):  1.2 
mi.  E Landa  de  Matamoros  ( TCWC  2969 1 );  1.5  mi.  E Landa 

Bezy:  Systematics  of  Lepidophyma  in  Mexico 


]J 


(TCWC  33063).  SAN  LUIS  POTOSI:  (17):  Boa  Capulin 
(LSUMZ  2379-80). 

L.  sylvaticum 

HIDALGO:  (11):  5.8  mi.  (by  Hwy  105)  S Tianquistengo 
(LACM  106741-48);  4.0  mi.  (by  Hwy  105)  S Tianquistengo 
(LACM  106721);  3 mi.  S Tianquistengo  (UIMNH  26230). 
NUEVO  LEON:  (1):  La  Boca  (KU  92612-13);  ca.  7 km  NE 
Santiago,  Presa  La  Boca  (LACM  106781-792);  (2):  5 mi.  N 
Las  Ajuntas  (EAL  4644).  QUERETARO:  ( 1 0):  El  Madrono, 
3.5  mi.  W (rd)  El  Lobo  [and  vie]  (LACM  109771,  SAM  1 104, 
TCWC  29692-29707,  32291,  33064,  35607,  UMMZ 
129749).  SAN  LUIS  POTOSI:  (3):  Buenavista  (ca.  20  mi. 
NE  Cerritos)  (AMNH  64025);  (4):  Alvarez  (58  kilo)  (MCZ 
24507-08);  between  San  Francisco  and  Alvarez  (MCZ 
157826);  Valle  de  los  Fantasmos  (SDNHM  60482);  (8):  27 
km  (by  Mex  80)  W El  Naranjo  (LACM  1 3 1 145-48);  3.8  mi. 
(by  Hwy  80)  NNE  Ciudad  del  Maiz  (LACM  131144);  5 mi. 
NE  Ciudad  del  Maiz  (TCWC  35582);  (9):  Huichihuagan 
(FMNH  39631).  TAMAULIPAS:  (5):  8 mi.  S,  6 mi.  W Vic- 
torio.  Sierra  Madre  Oriental  (KU  33992);  (6):  Rancho  del 
Cielo  [and  vie]  (AMNH  107273,  LACM  106751-60, 
106762-65,  LSUMZ  10989,  UMMZ  101301,  101375, 
102977-81,  109763-67);  (7):  Sierra  de  Tamaulipas,  Santa 
Maria  (UMMZ  102889-90);  10  mi.  W,  2 mi.  S Piedra  (KU 
33993-94).  VERACRUZ:  (12):  4 km  W Tlapacoyan  (KU 
26909). 

L.  flavimaculatum 

CHIAPAS:  (22):  Palenque  (LACM  65117-19);  Ruinas  de 
Palenque  (EAL  3030-31,  FSM  32915-16,  KU  94104-05); 
San  Juanito,  Palenque  (USNM  1 1 1486-87);  (23):  4.5  km  S 
Pichucalco  (KU  94106);  (24):  El  Estoracan,  ca.  50  km  N 
Cintalapa  (AMNH  73468);  (25):  Chiapa,  1 mi.  W (TNHC 
27517-18);  (26):  Ocozocoautla  Selvas  El  Ocote  (MCZ  5432 1— 
22);  16.1  mi.  NW  Ocozocoautla  (LACM  61259);  26  km  N 
Ocozocoautla  (UTEP  5367-68);  32  km  NW  Ocozocoautla, 
Selva  del  Ocote  (JFC);  25  mi.  (by  rd  to  Malpaso)  NW  Oco- 
zocoautla (UAZ  28764,  28805-07);  12  km  N Berriozabal 
(UTEP  5365-66);  (27):  ca.  5 km  S Solusuchiapa  (UAZ  31635); 
(28):  Lago  Miramar,  near  San  Quintin  (JFC);  (29):  Lacanja 
(LACM  114244).  OAXACA:  (30):  2.8  mi.  N Rio  Sarabia 
(UMMZ  1 15096);  (31):  2 km  S Tolosita  (KU  39676);  (32): 
Rio  Mono  Blanco,  Juchitan  (UIMNH  36832);  (33):  La  Gloria 
(UIMNH  35515);  (34):  Mogone  (UIMNH  40811);  (35):  50.5 
mi.  S Acayucan,  Hwy  185  (TNHC  25182).  QUINTANA 
ROO:  (36):  4. 1 km  NE  Felipe  Carrillo  Puerto  (UMRC  79- 
252).  TABASCO:  (37):  Teapa  (LACM  61260-61,  LSUMZ 
6878-79,  UIMNH  47883,  UMMZ  1 13777);  (38):  Soledad 
(UIMNH  47884).  VERACRUZ:  (39):  20  km  E Jesus  Car- 
ranza (KU  24453);  25  km  SE  Jesus  Carranza  (KU  26920- 
21);  35  km  SW  Jesus  Carranza  (KU  26919);  (40):  Rio  de  las 
Playas  (USNM  118638). 

L.  pajapanensis 

VERACRUZ;  (41):  Sontecomapan,  Los  Tuxtlas  [and  vie] 
(TCWC  21365,  UAZ  28765,  28808-1 1,  UTAR  3107,  31 10, 


3116,  TCWC  21365);  Coyame,  9 mi.  (by  rd)  SE  Catemaco 
(UAZ  28804);  Univ.  Mex.  Biol.  Exp.  Sta.,  ca.  33  km  ENE 
Catemaco  (TCWC  53351);  Coloma  de  Bastonal,  above  Que- 
zalapam  (TCWC  19133);  Laguna  Catemaco,  nr  Cuezalapan 
(UMMZ  126363-64);  4 mi.  SETebanca,  Los  Tuxtlas  (TCWC 
21364);  S slope  Volcan  San  Martin  (KU  97290,  UMMZ 
1 18220,  126362);  (42):  35  km  SE  Jesus  Carranza  (KU  269 1 3). 

L.  smithii 

CHIAPAS:  (43):  La  Esperanza  (UIMNH  10952-56,  10958- 
59,  10963,  10965,  10968-69,  10970-71,  10975-79,  10997- 
98);  (44);  Tonala  (UIMNH  26227-29).  GUERRERO:  (45): 
2 km  W Puerto  Marquez  (CU  9676-79,  9692-93,  9772, 
LACM  128590,  130027-29). 

L.  tarascae 

MICHOACAN:  (46):  near  Mexiquillo,  Aquila  District  (ENCB 
9221-22,  LACM  134226). 

L.  tuxtlae 

CHIAPAS:  (47):  25  mi.  (by  rd  to  Malpaso)  NW  Ocozocoautla 
(UAZ  28780-82).  OAXACA:  (48):  Vista  Hermosa  (KU 
87396-98);  30  mi.  (by  rd)  NE  Llano  de  las  Flores  (UMMZ 
125870);  (49):  Mts  nr  La  Gloria  (UIMNH  37236);  (50):  Finca 
San  Carlos,  Matias  Romero  Oaxaca  (FSM  32918).  VERA- 
CRUZ: (51):  Volcan  San  Martin  (TCWC  22102-03,  TNHC 
29792-93,  UIMNH  80695-99,  UMMZ  1 18219,  121165, 
122112,  126360-61);  S slope  Volcan  San  Martin  Tuxtla  (KU 
59560);  Rancho  El  Tular,  1 5 mi.  N San  Andres  Tuxtla  (USNM 
139731);  Rio  Tecolapan,  2.4  mi.  NNW  Tapalapan  (UMMZ 
115098-99);  Salto  de  Eyipantla  (TCWC  19134);  Montepio 
(FSM  32917);  Sontecomapan  [and  vie]  (CM  41470,  FSM 
32914,  TCWC  19135,  26717,  UAZ  28770-79,  UTAR  3101, 
3103-04,  3108-09,  31  1 1-13,  3115,  3127);  7.7  mi.  NW  Son- 
tecomapan (UTAR  3728-30,  3733-34);  E of  Lago  Catemaco, 
1 2.7  mi.  from  Catemaco  by  rd  (LACM  106795);  1 8 mi.  NNE 
Catemaco  (JCL  67);  Univ.  Mex.  Bio.  Exp.  Sta.,  ca.  33  km 
ENE  Catemaco  (TCWC  53352—53);  between  Laguna  Cate- 
maco and  Volcan  Martin  (UMMZ  121 166);  Coyame  (UAZ 
28763);  midway  between  Coyame  and  Tebanca  (UMMZ 
121164);  4 mi.  SE  Tebanca  (TCWC  21366);  5.6  mi.  ESE 
Tebanca  (UTAR  3156);  Rio  Quetzalapan  [and  vie]  (TCWC 
19136,  21367-69,  UTAR  3133,  3139-40);  Coloma  de  Bas- 
tonal (TCWC  19137);  Dos  Arroyos,  5 mi.  E Zapoapan  (TCWC 
21370-71);  (52):  25  km  SE  Jesus  Carranza  (KU  26912). 


ACKNOWLEDGMENTS 

I thank  the  following  individuals  for  assistance  with  aspects 
of  the  study:  Kathryn  Bolles,  Salvador  Contreras  B.,  Robb 
Dean,  James  R.  Dixon,  Jerry  D.  Johnson,  Carl  S.  Lieb,  Rob- 
ert I.  Lonard.  James  F.  Lynch,  Barbara  T.  Warburton,  and 
John  W.  Wright  for  held  assistance  and/or  collecting  live 
material;  Ticul  Alvarez  and  officials  of  the  Fauna  del  Silvestre 
for  granting  the  necessary  collecting  permits;  John  P.  Karges 
and  John  W.  Wright  for  discussions,  advice,  and/or  review 
of  the  manuscript;  John  DeLeon  for  photography;  Kathryn 


Contributions  in  Science,  Number  349 


Bezy:  Systematics  of  Lepidophyma  in  Mexico  15 


Bolles  for  illustration;  Beatriz  Larrain  for  translation;  Terri 
Togiai  for  typing  the  manuscript;  and  C.J.  Cole  (AMNH), 
B.C.  Brown  (BCB),  A.E.  Leviton  (CAS),  C.J.  McCoy  (CM), 
J.M.  Savage  (CRE),  F.H.  Pough  (CU),  E.A.  Liner  (EAL),  T. 
Alvarez  (ENCB),  R.F.  Inger,  H.  Marx,  ELK.  Voris  (FMNH), 
W.  Auffenberg  (FSM),  W.E.  Duellman  (KU),  J.F.  Copp  (JFC), 
D.A.  Rossman  (LSUMZ),  P.  Alberch,  E.E.  Williams  (MCZ), 
FEW.  Greene,  D.B.  Wake  (MVZ),  T.M.  Uzzell  (PAS),  S.A. 
Minton  (SAM),  G.K.  Pregill  (SDNHM),  J.R.  Dixon  (TCWC), 
R.F.  Martin  (TNHC),  C.H.  Lowe  (UAZ),  D.F.  Hoffmeister 
(UIMNH),  A.G.  Kluge  (UMMZ),  J.C.  Lee  (UMRC),  G.R. 
Zug  (USNM),  W.F.  Pyburn  (UTA),  and  R.G.  Webb  (UTEP) 
for  loan  of  specimens  under  their  charge  (museum  abbre- 
viations follow  Leviton  et  al.,  1980). 

LITERATURE  CITED 

A vise,  J.C.,  and  R.K.  Selander.  1972.  Evolutionary  genetics 
of  cave-dwelling  fishes  of  the  genus  Astyanax.  Evolution 
26(1):  1-19. 

Bezy,  R.L.  1972.  Karyotypic  variation  and  evolution  of  the 
lizards  in  the  family  Xantusiidae.  Contributions  in  Sci- 
ence 227;  1-29. 

. 1973.  A new  species  of  the  genus  Lepidophyma 

(Reptilia:  Xantusiidae)  from  Guatemala.  Contributions 
in  Science  239: 1-7. 

Bezy,  R.L.,  R.G.  Webb,  and  T.  Alvarez.  1982.  A new  species 
of  the  genus  Lepidophyma  (Sauria:  Xantusiidae)  from 
Michoacan,  Mexico.  Herpetologica  38(3):36 1-366. 
Blackburn,  D.G.  1982.  Evolutionary  origins  of  viviparity 
in  the  Reptilia.  I.  Sauria.  Amphibia-Reptilia  3(2/3):  185- 
205. 

Bocourt,  M.  1876.  Note  sur  quelques  reptiles  de  l’isthme 
de  Tehuantepec  (Mexique)  donnes  par  M.  Sumichrast 
au  Museum.  Journal  de  Zoologie,  Paris  5:386-41  1. 
Bull.J.J.  1980.  Sex  determination  in  reptiles.  The  Quarterly 
Review  of  Biology  55(1): 3—2 1 . 

Darevsky,  I.S.,  L.A.  Kupriyanova, and  M.A.  Bakradze.  1 978. 
Occasional  males  and  intersexes  in  parthenogenetic 
species  of  Caucasian  rock  lizards  (genus  Lacerta).  Copeia 
1978(2):201-207. 

Dixon,  J.R.,  C.A.  Ketchersid.  and  C.S.  Lieb.  1972.  The 
herpetofauna  of  Queretaro,  Mexico,  with  remarks  on 
taxonomic  problems.  Southwestern  Naturalist  16(3  & 
4):225-237. 

Dixon,  W.J.  [Editor],  1981.  BMDP  Statistical  software. 

University  of  California  Press,  Berkeley. 

Dumeril,  A.M.C.,  and  A.H.A.  Dumeril.  1851.  Catalogue 
methodique  de  la  collection  des  reptiles.  Museum  d’His- 
toire  Naturelle  de  Paris. 

Hall,  W.P.,  and  R.K.  Selander.  1973.  Hybridization  of 


karyotypically  differentiated  populations  in  the  Scelo- 
porus  grammicus  complex  (Iguanidae).  Evolution  27(2): 
226-242. 

Leviton,  A.E.,  R.  McDiarmid,  S. Moody,  M.  Nickerson,  J. 
Rosado,  O.  Sokol,  and  H.  Voris.  1980.  Museum  ac- 
ronyms—second  edition.  Herpetological  Review  11(4): 
93-102. 

Leopold,  A.S.  1959.  Wildlife  of  Mexico.  The  game  birds 
and  mammals.  University  of  California  Press,  Berkeley. 

Mitchell,  R.W.,  W.H.  Russell,  and  W.R.  Elliott.  1977. 
Mexican  eyeless  characin  fishes,  genus  Astyanax : envi- 
ronment, distribution,  and  evolution.  Special  Publica- 
tions, The  Museum,  Texas  Tech  University  12:1-89. 

Mosauer,  W.  1936.  A new  xantusiid  lizard  of  the  genus 
Lepidophyma.  Herpetologica  1(1  ):3— 5 + Plate  II. 

Savage,  J.M.  1963.  Studies  on  the  lizard  family  Xantusiidae 
IV.  The  genera.  Los  Angeles  County  Museum  Contri- 
butions in  Science  71:1-38. 

Simpson,  G.G.,  A.  Roe,  and  R.C.  Lewontin.  1960.  Quan- 
titative zoology.  Revised  Edition.  Harcourt,  Brace,  and 
Co.,  New  York. 

Sites,  J.W.  1983.  Chromosome  evolution  in  the  iguanid 
lizard  Sce/oporus  grammicus.  I.  Chromosome  polymor- 
phisms. Evolution  37(  1 ):38— 53. 

Smith,  H.M.  1939.  Notes  on  Mexican  reptiles  and  am- 
phibians. Zoological  Series  of  Field  Museum  of  Natural 
History  24(4):  15-35. 

. 1942.  Mexican  herpetological  miscellany.  Proceed- 
ings of  the  United  States  National  Museum  92(3153): 
349-395. 

. 1973.  A tentative  rearrangement  of  the  lizards  of 

the  genus  Lepidophyma.  Journal  of  Herpetology  7(2): 
109-123. 

Smith,  H.M.,  and  M.  Alvarez  del  Toro.  1977.  A new  trog- 
lodytic  lizard  (Reptilia,  Lacertilia,  Xantusiidae)  from 
Mexico.  Journal  of  Herpetology  1 1(1): 37—40. 

Taylor,  E.H.  1939.  A new  species  of  the  lizard  genus  Lep- 
idophyma from  Mexico.  Copeia  1939(3):  13 1-1 33. 

Walker,  C.F.  1955.  Two  new  lizards  of  the  genus  Lepido- 
phyma from  Tamaulipas.  Occasional  Papers  of  the  Mu- 
seum of  Zoology  University  of  Michigan  564:1-10. 

Werler,  J.E.  1957.  A new  lizard  of  the  genus  Lepidophyma 
from  Volcan  San  Martin  Pajapan,  Veracruz.  Herpeto- 
logica 1 3(3):223— 226. 

Werler,  J.E. , and  F.A.  Shannon.  1957.  A new  lizard  of  the 
genus  Lepidophyma  from  Veracruz,  Mexico.  Herpeto- 
logica 13(2):  1 19-122. 

White,  M.J.D.  1973.  Animal  cytology  and  evolution.  Third 
Edition.  Cambridge  University  Press,  London. 

Submitted  16  May  1983;  accepted  13  September  1983. 


16  Contributions  in  Science,  Number  349 


Bezy:  Systematics  of  Lepidophyma  in  Mexico 


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MEGAPALEONTOLOGY  OF  THE  EOCENE  LLAJAS  FORMATION, 
SIMI  VALLEY,  CALIFORNIA 

Richard  L.  Squires 


Contributions  in  Science,  Number  350 
Natural  History  Museum  of  Los  Angeles  County 
13  July  1984 


ISSN  0459-8113 


Natural  History  Museum  of  Los  Angeles  County 
900  Exposition  Boulevard 
Los  Angeles,  California  90007 


CONTENTS 


ABSTRACT  1 

INTRODUCTION  1 

HISTORICAL  REVIEW  2 

MEGAFOSSILS 2 

DEPOSITIONAL  ENVIRONMENTS  AND  STRATIGRAPHIC  DISTRIBUTION 

OF  MEGA  FOSSILS  4 

PAI.EOCL.IMATE 7 

AGE  9 

CORRELATION 1 1 

BIOGEOGRAPHY 1 1 

SYSTEMATIC  MATERIALS  AND  METHODS 13 

SYSTEMATICS 13 

Phylum  Protista 13 

Order  Foraminiferida  13 

Family  Discocyclinidae 13 

Pseudophragmina  ( Proporocyclina ) clarki  (Cushman,  1920) 14 

Phylum  Coelenterata 14 

Order  Scleractinia 14 

Family  Caryophylliidae 14 

ITrochocyathus  striatus  (Gabb,  1864)  14 

Turbino/ia  dickersoni  Nomland,  1916 15 

Phylum  Brachiopoda 15 

Order  Terebratulida 1 5 

Eogryphus  tolmani  Hertlein  and  Grant,  1944  15 

Phylum  Annelida 1 5 

Order  Sedentaria 1 5 

Family  Serpulidae  15 

Rotularia  tejonense  ( Arnold,  1910) 15 

Phylum  Mollusca 15 

Class  Scaphopoda 15 

Family  Dentaliidae 15 

Dentalium  stentor  Anderson  and  Hanna,  1925  16 

Dentalium  (Laevidentalium)  calafium  Vokes,  1939  16 

Class  Gastropoda 16 

Order  Archaeogastropoda 16 

Family  Neritidae  16 

Nerita  cf.  N.  ( Amphinerita ) eorex  Vokes,  1939  16 

Velates  perversus  ( Gmelin,  1791)  16 

Order  Mesogastropoda  17 

Family  Turritellidae 17 

Turrit ella  meganosensis  protumescens  Merriam  and  Turner,  1937  17 

Turrite/la  andersoni  Dickerson,  1916 18 

Turritella  andersoni  lawsoni  Dickerson,  1916  18 

Turritella  buwa/dana  Dickerson,  1916 18 

Turritella  uvasana  infera  Merriam,  1941  19 

Turritella  uvasana  applinae  Hanna,  1927  19 

Family  Architectonicidae 19 

Architectonica  ( Architectonica ) l/ajasensis  Sutherland,  1966  19 

Architectonica  ( Stellaxis ) cognata  Gabb,  1864  19 

Family  Cerithiidae 20 

Benoistia  umpquaensis  Turner,  1938  20 

Family  Cerithiopsidae 20 

Cerithiopsis  llajasensis  n.  sp 20 

Family  Epitoniidae 21 

Cirsotrema  sp 21 


Family  Calyptraeidae 21 

Calyptraea  diegoana  (C onrad,  1855) 21 

Family  Xenophoridae 21 

Xenophora  stocki  Dickerson,  1916 21 

Family  Strombidae 21 

Ectinochilus  (Macilentos)  macilentus  (White,  1889) 21 

Family  Seraphsidae 23 

Paraseraphs  erraticus  (C ooper,  1894) 23 

Family  Cypraeidae 23 

Cypraea  castacensis  Stewart,  1927  23 

Family  Naticidae 24 

Eocernina  hannibali  Dickerson,  1914 24 

Tejonia  moragai  (Stewart,  1927) 24 

Pachycrommium  clarki  (Stewart,  1927) 24 

Polinices  (Euspira)  nuciformis  (Gabb,  1864)  25 

Neverita  (Neverita)  globosa  Gabb,  1 869  25 

Sinum  obliquum  (Gabb,  1864)  26 

Natica  ( Naticarius ) aff.  N.  (TV.)  uvasana  Gabb,  1864  26 

Family  Cassididae  26 

Galeodea  ( Cahagaleodea)  californica  Clark,  1942  26 

Galeodea  (Gomphopages)  susanae  Schenck,  1926  27 

Phalium  (Semicassis)  tuberculiformis  ( Hanna,  1924) 27 

Family  Cymatiidae 27 

Cymatium  (Septa)  janetae  Squires,  1983a  27 

Ranella  katherineae  Squires,  1983a 29 

Family  Bursidae 29 

Olequahia  domenginica  (Vokes,  1939) 29 

Rane/lina  pilshryi  Stewart.  1927  29 

Family  Ficidae  29 

Ficopsis  cooperiana  Stewart,  1927  29 

Ficopsis  remondii  crescentensis  Weaver  and  Palmer,  1922  30 

Order  Neogastropoda  30 

Family  Muricidae 30 

Laevityphis  (Laevityphis)  antiquus  (Gabb,  1864) 30 

Family  Buccinidae 30 

A ncistrolepisl  carolineae  n.  sp 30 

Family  Nassariidae  31 

Molopophorus  cretaceus  (Gabb,  1864) 31 

Family  Fasciolariidae  31 

Clavilithes  tabulatus  (Dickerson,  1913) 31 

Clavilithes  n.  sp.  A Clark  and  Vokes,  1936  31 

Clavilithes  n.  sp.  B Squires,  1983a 31 

Fusinus  teglandae  Hanna,  1927  33 

Fusinus  aff.  F.  ucalius  X okes,  1939  33 

Family  Olividae 33 

Pseudoliva  lineata  Gabb,  1864  33 

Strepsidura  ficus  (Gabb,  1864) 33 

Ancilla  (Spirancilla)  gabbi  Cossmann,  1899  34 

Olive/ 1 a mathewsonii  Gabb,  1864  34 

Family  Mitridae 34 

Proximitra ? cretacea  (Gabb,  1864) 34 

Family  Tudiclidae  34 

PseudoperissolaxblakeipraeblakeiXoV.es, , 1939  34 

Family  Harpidae 35 

Eocithara  mutica  californiensis  (Vokes,  1937) 35 

Family  Voiutidae 35 

Cryptochorda  (Cryptochorda)  californica  (Cooper,  1894) 35 


Lyria  andersoni  Waring,  1917 35 

Lyrischapa  lajollaensis  ( Hanna,  1927) 36 

Family  Cancellariidae 36 

Bonellitia  (Admetula)  paucivaricata  (Gabb,  1864) 36 

Family  Turridae 36 

Pleurofusia  fresnoensis  (Arnold,  1910) 36 

Fusiturricula  ( Crenaturricula ) crenatospira  (Cooper,  1894) 36 

Fusiturricula  ( Crenaturricula ) crenatospira  domenginica  Vokes,  1939  . . 37 

Surculites  mathewsonii  (Gabb,  1864) 37 

Domenginelia  claytonensis  ( Gabb,  1864) 37 

Fxilia  ilajasensis  Bentson,  1940  39 

Family  Conidae  39 

Conus  caleocius  \ okes,  1939  39 

Conus  hornii  umpquaensis  Turner,  1938  39 

Family  Terebridae 39 

Terebra  calij arnica  Gabb,  1869  39 

Order  Cephalaspidea 39 

Family  Cylichnidae 39 

Cylichnina  tantilla  (Anderson  and  Hanna,  1925) 40 

Scaphander  (Mirascapha)  costatus  (Gabb,  1864) 40 

Family  Philinidae 40 

Megistostoma  gabbianum  ( Stoliczka,  1868) 40 

Class  Bivalvia 40 

Order  Nuculoida 40 

Family  Nuculidae 40 

Acila  (Truncaci/a)  decisa  (Conrad.  1855)  41 

Family  Nuculanidae  41 

Nuculana  (Saccella)  gabbii  (Gabb,  1864)  41 

Order  Arcoida 41 

Family  Glycymerididae  41 

Glycymeris  (Glycymeris)  roseca nyonensis  Hanna,  1927  41 

Glycymeris  (Glycymerita)  sagittata  ( Gabb,  1864) 42 

Order  Mytiloida 42 

Family  Mytilidae 42 

Brachidontes  (Brachidontes)  cowlitzensis  (Weaver  and  Palmer,  1922)  ..  42 

Family  Pinnidae 42 

Pinna  lewisi  Waring,  1917 42 

Pinna  Ilajasensis  Squires,  1983a 43 

Order  Pterioida 43 

Family  Malleidae 43 

Nayadina  (Exputens)  Ilajasensis  (Clark,  1934) 43 

Family  Spondylidae 43 

Spondylus  carlosensis  Anderson,  1905  43 

Family  Anomiidae 45 

Anomia  mcgoniglensis  Hanna,  1927  45 

Family  Ostreidae  45 

Ostrea  idriaensis  Gabb,  1869  45 

Order  Veneroida 45 

Family  Lucinidae 45 

Claibornites  diegoensis  { Dickerson,  1916) 45 

Family  Carditidae  46 

Venericardia  (Pacifcor)  hornu  calafia  Stewart,  1930  46 

Venericardia  ( Pacificor ) aragonia  joaquinensis  (Vokes,  1939) 46 

Glyptoactis  (G/yptoactis)  domenginica  (V okes,  1939) 46 

Family  Crassatellidae  47 

Crassatella  uvasana  Conrad,  1855  47 

Family  Cardiidae 49 


Acanthocardia  (Schedocardia)  breweru  (Gabb,  1864) 49 

Nemocardium  linteum  (Conrad,  1855)  49 

Family  Solenidae 50 

Solena  (Eosolen)  novacularis  (Anderson  and  Hanna,  1928)  50 

Family  Tellinidae 50 

Macoma  rosa  Hanna,  1927  50 

Family  Psammobiidae  50 

Gari  cf.  G.  eoundulata  Yokes.  1939  50 

Family  Veneridae 50 

Callista  ( Costacallista ) cf.  C.  (C.)  hornii  (Gabb,  1864)  51 

Callocardia  (Nitidavenus)  tejonensis  (Waring,  1914)  51 

Pitar  (Calpitaria)  uvasanus  ( Conrad,  1855)  51 

Pitar  (Lamelliconcha)  joaquinensis  Vokes,  1939  51 

Marcia  {Mercimonia)  bunkeri  (Hanna,  1927)  53 

Order  Myoida 53 

Family  Corbulidae 53 

Corbula  (Caryocorbula)  dickersoni  ( Weaver  and  Palmer,  1 922)  53 

Family  Teredinidae 53 

Teredo ? sp 53 

Order  Pholadomyoida 54 

Family  C’uspidariidae  54 

Cardiomya  aff  C.  russelli  (Hanna,  1927) 54 

Class  Cephalopoda 54 

Order  Nautilida  54 

Family  Aturiidae  54 

Aturia  myrlae  Hanna,  1927  54 

Order  Sepiida  55 

Family  Indeterminate 55 

Spirulimorph  sepiid 55 

Phylum  Arthropoda 55 

Infraorder  Brachyura 55 

Family  Goneplacidae 55 

Glyphithyreus  weaveri  (Rathbun,  1926) 55 

Phylum  Kehinodermata  55 

Order  Spatangoida 56 

Family  Schizasteridae 56 

Schizaster  diabloensis  Kew,  1920  56 

Phylum  Chordata 56 

Order  Lamniformes 56 

Family  Odontaspididae  56 

Odontaspis  sp 56 

Family  Lamnidae 57 

Isurus  cf.  /.  praecursor  (Leriche,  1906) 57 

LOCALITIES 57 

ACKNOWLEDGMENTS  66 

LITERATURE  CITED 67 


MEGAPALEONTOLOGY  OF  THE  EOCENE  LLAJAS  FORMATION 
SIMI  VALLEY,  CALIFORNIA 


Richard  L.  Squires1 


ABSTRACT.  The  paleontology  and  stratigraphic  distribution  of  the 
megafossils  in  the  late  early  through  early  middle  Eocene  age  Llajas 
Formation,  Simi  Valley,  Los  Angeles  and  Ventura  Counties,  south- 
ern California,  are  described  in  detail.  One  hundred  and  seven  species 
and  subspecies,  representing  88  identifiable  genera,  are  recorded  for 
132  localities.  Twenty-six  of  these  species  are  previously  unreported 
from  the  Llajas  Formation. 

Illustrations,  synonymies,  primary  type  material  information.  West 
Coast  molluscan  stage  ranges,  geographic  distributions,  local  occur- 
rences, and  remarks  are  provided  for  the  taxa,  which  include  one 
large  foraminifer,  two  corals,  one  brachiopod,  one  annelid,  two 
scaphopods,  65  gastropods,  29  bivalves,  one  nautiloid,  one  sepiid, 
one  brachyuran,  one  spatangoid,  and  two  sharks.  The  sepiid  can  be 
identified  only  to  the  family  level.  A new  species  of  Cerithiopsis  and 
a new  species  of  Ancistrolepis!  are  named.  Crassatella  uvasana  and 
C.  u.  semidentata  are  shown  to  be  identical. 

Most  of  the  megafossils  occur  in  channel-fill  deposits  in  shallow- 
marine  strata  but  have  undergone  minimal  postmortem  transport. 
The  Llajas  megafauna  lived  in  warm  waters  less  than  80  m depth. 

Mollusks  indicative  of  the  lower  of  the  two  faunal  zones  of  the 
West  Coast  provincial  molluscan  “Capay  Stage”  (lower  Eocene)  are 
present  in  the  lowermost  fossil-bearing  beds  of  the  Llajas  Formation. 
The  remaining  fossil-bearing  part  of  the  Llajas  contains  mollusks 
indicative  of  the  “Domengine  Stage”  (upper  lower  through  lower 
middle  Eocene).  Age  refinement  of  the  Llajas  extends  the  molluscan 
stage  ranges  of  several  molluscan  species.  The  Llajas  megafauna  is 
similar  to  those  in  late  early-early  middle  Eocene  age  formations 
from  San  Diego,  California  through  southwestern  Oregon.  These 
megafaunas  were  in  the  same  faunal  province  that  extended,  at  least, 
from  southern  California  into  Washington.  Most  of  the  genera  were 
already  present  on  the  West  Coast  by  Llajas  time  and  several  species 
were  carry-overs  from  earlier  times.  A few  genera  were  early  Eocene 
immigrants  from  the  Caribbean  and  Old  World  Tethyan  regions, 
and  some  were  early  middle  Eocene  immigrants  from  the  same 
regions.  Several  genera  originated  on  the  West  Coast  during  the  early 
middle  Eocene. 

INTRODUCTION 

Since  the  early  1900’s,  paleontologists  have  collected  mega- 
fossils from  the  late  early  through  early  middle  age  Eocene 
Llajas  Formation  in  the  Simi  Valley  area,  Los  Angeles  and 

Contributions  in  Science,  Number  350,  pp.  1-76 
Natural  History  Museum  of  Los  Angeles  County,  1984 


Ventura  Counties,  southern  California.  The  fossils  are  pre- 
dominantly mollusks  and  nearly  every  paleontological  mu- 
seum on  the  West  Coast  has  a representative  collection.  To 
date,  however,  there  has  been  no  comprehensive  taxonomic 
and  stratigraphic  analysis  of  the  Llajas  megafauna.  Research- 
ers, therefore,  have  been  hindered  in  performing  detailed 
comparative  studies  on  the  taxonomy,  biogeography,  and 
time-correlation  of  the  Llajas  megafossils.  The  objectives  of 
this  paper  are  1)  to  tabulate  the  species  and  show  their  dis- 
tribution within  the  formation,  2)  to  provide  synonymies 
and  illustrations  of  the  species,  3)  to  provide  taxonomic  re- 
finements of  certain  known  taxa  and  describe  two  new  gas- 
tropods, 4)  to  interpret  the  paleoenvironment  and  biogeog- 
raphy of  the  megafauna,  5)  to  refine  the  age  assignment  and 
correlation  of  the  Llajas  Formation,  and  6)  to  refine  the  West 
Coast  molluscan  stage  ranges  of  the  taxa. 

The  Llajas  Formation  crops  out  in  the  southwestern  Santa 
Susana  Mountains  and  along  the  south  side  of  Simi  Valley 
(Fig.  1 ).  It  disconformably  overlies  the  late  Paleocene  through 
earliest  Eocene  age  marine  Santa  Susana  Formation  and, 
except  where  local  faults  occur,  is  unconformably  overlain 
by  the  early  late  Eocene  through  Oligocene  age  nonmarine 
Sespe  Formation  (Figs.  2 and  3). 

Outcrops  of  the  Llajas  Formation  are  typically  extensively 
covered.  Best  exposures  are  at  the  545-m-thick  type  section 
in  the  southwestern  Santa  Susana  Mountains  (Fig.  1 ).  In  1981 
and  1 983b,  I reported  that  the  Llajas  Formation  in  the  south- 
western Santa  Susana  Mountains  consists  mostly  of  a trans- 
gressive (retrogradational)  sequence  of  facies  that  grades  ver- 
tically from  coastal  alluvial  fan,  to  shallow  marine,  to  outer 
shelf  and  slope  with  incised  channels.  The  outer  shelf  and 
slope  facies  was  subsequently  covered  by  regressive  (progra- 
dational)  shallow-marine  facies.  The  regressive  sequence  is 


1.  Department  of  Geological  Sciences,  California  State  Univer- 
sity, Northridge,  California  91330,  and  Research  Associate,  Inver- 
tebrate Paleontology,  Natural  History  Museum  of  Los  Angeles  Coun- 
ty. 


ISSN  0459-8113 


118°  40' 


incomplete  due  to  erosional  truncation  by  the  Sespe  For- 
mation (Fig.  2). 

HISTORICAL  REVIEW 

In  the  last  70  years,  starting  with  Waring  (1914),  there  have 
been  many  investigations  dealing  with  the  megafossils  of  the 
Llajas  Formation.  These  previous  studies  have  included  par- 
tial faunal  lists  (Waring,  1917;  Clark,  1926;  Hanna,  1927; 
McMasters,  1932;  Cushman  and  McMasters,  1936;  Turner, 
1938;  Vokes,  1939;  Lewis,  1940;  Seiden,  1972),  descriptions 
of  new  taxa  (Waring,  1914,  1917;  Schenck,  1926;  Stewart, 
1927,  1930;  Clark,  1934,  1942;  Merriam  and  Turner,  1937; 
Vokes,  1937,  1939;  Bentson,  1940;  Merriam,  1941;  Hertlein 
and  Grant,  1944;  Sutherland,  1966;  Squires,  1983a),  and 
taxonomic  refinements  or  miscellaneous  comments  on  known 
species.  In  this  last  category,  the  most  notable  papers  are 
those  of  Schenck  (1931),  Gardner  and  Bowles  (1934),  Clark 
and  Vokes  (1936),  Turner  (1938),  Vokes  (1935,  1939),  Mer- 
riam (1941),  Hanna  and  Hertlein  (1943),  Verastegui  (1953), 
Marincovich  (1977),  Givens  (1979),  Squires  (1979),  and  Saul 
(1983). 


As  the  name  “Llajas  Formation”  was  not  formalized  until 
1936  by  Cushman  and  McMasters,  earlier  workers  referred 
to  such  strata  by  various  names  (Fig.  4).  Clark  and  Vokes 
(1936),  Turner  (1938),  and  Vokes  (1939)  used  the  terms 
“Lower  Llajas”  and  “Upper  Llajas”  but  did  not  define  them. 

Various  aspects  of  the  microfossils  of  the  Llajas  Formation 
have  been  reported  on  by  McMasters  (1932),  Cushman  and 
McMasters  (1936),  Laiming  ( 1 940a,  1940b,  1943),  Mallory 
(1959),  Schymiczek  and  Squires  (1981),  Schymiczek  (1983a, 
1983b),  and  Filewicz  and  Hill  (1983). 

MEGAFOSSILS 

One  hundred  and  seven  taxa,  more  than  90  percent  of  which 
are  mollusks,  were  identified  from  the  Llajas  Formation. 
Taxa  identified  to  species  and  subspecies  include  64  gastro- 
pods, 28  bivalves,  two  scaphopods,  two  solitary  corals,  and 
one  species  each  of  large  foraminifer,  brachiopod,  annelid, 
nautiloid,  brachyuran,  and  spatangoid.  One  gastropod,  one 
bivalve,  and  two  sharks  are  identified  only  to  genus.  A spi- 
rulimorph  sepiid  could  only  be  identified  to  the  family  level. 
All  of  these  taxa  are  illustrated  in  Figures  5 through  1 3.  Other 


2 Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


NONMARINE 


Figure  2.  Stratigraphic  column  of  the  Llajas  Formation,  showing 
depositional  environments  based  on  Squires  (1981,  1983b). 


taxa,  too  poorly  preserved  for  even  generic  determination, 
include  a bryozoan,  two  brachiopods,  calcareous  worm  tubes, 
two  gastropods,  a nautiloid,  a myliobatoid,  and  scattered 
Teredo! -bored  wood  fragments. 

The  identifications  of  Llajas  species  and  subspecies  studied 
in  this  report  are  based  on  published  figures  and  descriptions 
and  selected  comparisons  with  type  specimens  and  non-type 
specimens  on  deposit  at  1)  Department  of  Earth  and  Space 
Sciences,  University  of  California  at  Los  Angeles,  2)  Uni- 
versity of  California,  Museum  of  Paleontology,  Berkeley,  3) 
Natural  History  Museum  of  Los  Angeles  County,  and  4) 
California  State  University,  Northridge. 

Megafossils  were  collected  at  121  localities  in  the  south- 
western Santa  Susana  Mountains  and  1 1 localities  along  the 
south  side  of  the  Simi  Valley.  All  of  the  localities  are  de- 
scribed in  the  “Localities”  section,  and  the  relative  strati- 


Figure  3.  Correlation  of  the  Llajas  Formation  with  Eocene  West 
Coast  molluscan  stages  (after  Saul,  1983);  millions  of  years  before 
present  (M.Y.B.P.)  scale,  epochs,  subepochs,  standard  ages,  planktic 
foraminifera  zones,  and  calcareous  nannoplankton  zones  (all  after 
Berggren,  Kent,  and  Flynn,  in  press);  and  West  Coast  benthic  fo- 
raminifera stages  (after  Poore,  1980). 


graphic  position  of  each  one  is  shown  in  Figures  14  through 
19. 

Species  and  subspecies  with  holotype  localities  in  the  Llajas 
Formation  are  listed  in  Table  1.  Species  not  previously  re- 
ported from  the  Llajas  Formation  are  listed  in  Table  2. 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  3 


Vyvvv^v/  Unconformity  Conformable  contact 

Nature  of  contact  not  discussed 

* Present  only  on  south  side  of  Simi  Valley 

Figure  4.  Comparative  concepts  of  Paleogene  stratigraphy  in  the 
Simi  Valley  area  (after  Squires,  1983b). 


Some  megafossil  species  previously  reported  (Waring,  1914, 
191 7;  Clark,  1921:158-159,  1926:1 14-1  16;  Clark,  1942;Kew, 
1924:25,  29;  Hanna,  1927:260;  Vokes,  1935,  1939:23-26, 
30-31;  Clark  and  Vokes,  1936;  Turner,  1938:33-37;  Bent- 
son,  1940;  Ingram,  1940)  from  the  Llajas  Formation  (or 
generally  equivalent  strata)  were  not  found  during  the  present 
study.  There  are  serious  problems  in  trying  to  substantiate 
the  records  of  these  species  because  1)  locality  information 
is  either  lacking  or  vague,  and/or  2)  previously  used  for- 
mation names  (i.e.,  Meganos  and  Domengine)  are  not  the 
exact  equivalents  of  the  Llajas  Formation  (Fig.  3).  In  addi- 
tion, because  these  species  names  usually  have  only  been 
listed,  without  illustration  or  reference  to  catalogued  museum 
specimens,  there  is  no  way  to  verify  the  identifications.  Some 
of  these  so-called  “Llajas”  species  can  be  shown  to  be  from 
the  underlying  late  Paleocene  through  earliest  Eocene  age 
Santa  Susana  Formation.  Based  on  analysis  of  Zinsmeister’s 
(1974,  1983a)  work  on  the  megafauna  of  the  Santa  Susana 
Formation,  the  following  species  are  from  this  unit:  Cucul- 
laria  morani  [=Cucullaea],  mentioned  by  Waring  (1914, 
1917);  Polinices  hornii,  Mesalia  martinezensis  [ = Turritella ], 
Turritella  infragranulata,  and  Septifer  elegans,  mentioned 
by  Waring  (1917).  Clark  (1942)  reported  that  the  type  locality 
of  Chedevi/lea  stewart i and  Tibia  (Eotibia)  Uajasensis  is  in 
the  lower  portion  of  the  Llajas  Formation.  According  to  Keen 
and  Bentson  (1944:143),  this  locality  (University  of  Califor- 
nia, Museum  of  Paleontology,  Berkeley,  locality  7015)  plots 
within  Paleocene  strata  as  mapped  by  Nelson  (1925). 

The  marine  strata  of  the  Llajas  Formation  have  been  thor- 
oughly bioturbated,  but  the  only  identifiable  trace  fossils  are 
Ophiomorpha,  Thalassinoides,  and  Chronditesl.  The  first  two 
ichnogenera  are  fairly  common  in  the  shallow-marine  facies, 
especially  in  certain  beds  where  this  facies  interfingers  with 
the  coastal  alluvial-fan  facies.  An  Ophiomorpha  burrow  in 
one  of  these  beds  is  figured  by  Squires  (1981:930,  fig.  6C; 


Table  1.  Megafossils  with  holotype  localities  in  the  Llajas  For- 
mation. 

Brachiopoda 

Eogryphus  tolmani  Hertlein  and  Grant,  1944 
Gastropoda 

A ncistrolepisl  carolineae  n.  sp. 

Architectonica  ( Architectonica ) Uajasensis  Sutherland,  1966 
Cerithiopsis  Uajasensis  n.  sp. 

Conus  caleocius  Vokes,  1939 
Cymatium  (Septa)  janetae  Squires,  1983 
Eocithara  mutica  californiensis  (Vokes,  1937) 

Exilia  Uajasensis  Bentson,  1940 

Fusiturricula  (Crenaturricula)  crenatospira  domenginica 
Vokes,  1939 

Galeodea  (Caliagaleodea)  californica  Clark,  1942 
Galeodea  ( Gomphopages ) susanae  Schenck,  1926 
Lyria  andersoni  Waring,  1917 
Pachycrommium  clarki  (Stewart,  1927) 

Ranella  katherineae  Squires,  1983 

Tectarius  ligniticus  Vokes,  1939  (see  Benoistia  umpquaensis) 
Turritella  andersoni  lawsoni  secondaria  Merriam,  1941 
Turritella  andersoni  susanae  Merriam,  1941 
Turritella  meganosensis  protumescens  Merriam  and 
Turner,  1937 

Turritella  uvasana  etheringtoni  Merriam,  1941 
(see  T.  u.  applinae) 

Turritella  uvasana  infera  Merriam,  1941 
Bivalvia 

Nayadina  (Exputens)  Uajasensis  (Clark,  1 934) 

Callocardia  (Nitidavenus)  tejonensis  (Waring,  1914) 

Pinna  lewisi  Waring,  1917 

Pinna  Uajasensis  Squires,  1983 

Venericardia  (Paciftcor)  hornii  calafia  Stewart,  1930 


1983b,  fig.  6E).  Chronditesl  is  common  in  the  outer  shelf 
and  slope  facies. 

DEPOSITIONAL  ENVIRONMENTS  AND 
STRATIGRAPHIC  DISTRIBUTION 
OF  MEGAFOSSILS 

Based  on  sedimentologic  and  lithologic  features  of  the  Llajas 
Formation,  on  comparative  studies  of  modem  and  ancient 
sedimentary  sequences,  and  ecology  of  representative  genera 
of  mollusks  and  benthic  foraminifers,  the  following  facies 
have  been  recognized  in  the  Llajas  Formation:  coastal  allu- 
vial fan,  shallow  marine,  and  outer  shelf  and  slope  with  in- 
cised channels.  In  a vertical  sense,  most  of  the  formation  is 
transgressive  (retrogradational)  in  that  shallower  facies  are 
overlain  by  deeper  facies.  The  uppermost  part  of  the  for- 
mation, however,  represents  a regressive  (progradational) 
event  in  which  the  outer  shelf  and  slope  facies  and  associated 


4 Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Table  2.  Megafossils  previously  unreported  from  the  Llajas  For- 
mation. 

Anthozoa 

ITrochocyathus  striatus  (Gabb,  1 864) 

Turbinolia  dickersoni  Nomland,  1916 

Annelida 

Rotularia  tejonense  (Arnold,  1909) 

Scaphopoda 

Dentalium  ( Laevidentalium ) calaftum  Vokes,  1939 
Dentalium  stentor  Anderson  and  Hanna,  1925 

Gastropoda 

Ancistrolepis ? carolineae  n.  sp. 

Cerithiopsis  llajasensis  n.  sp. 

Cirsotrema  sp. 

Laevityphis  ( Laevityphis ) antiquus  (Gabb,  1864) 

Molopophorus  cretaceus  (Gabb,  1864) 

Nerita  cf.  N.  (Amp  hi  writ  a)  eorex  Vokes,  1939 
Neverita  ( Neverita ) globosa  Gabb,  1 869 
Olivella  mathewsonii  Gabb,  1 864 
Terebra  California?  Gabb,  1869 

Bivalvia 

Anomia  mcgoniglensis  Hanna,  1927 
Cardiomya  aff.  C.  russet  1 1 (Hanna,  1927) 

Corbula  ( Caryocorbula)  dickersoni  Weaver  and  Palmer,  1922 

Gari  aff.  G.  eoundulata  Vokes,  1939 

Glycymeris  (Glycymeris)  rosecanyonensis  Hanna,  1927 

Macoma  rosa  Hanna,  1927 

Marcia  (Mercimonia)  bunkeri  (Hanna,  1927) 

Pilar  (Calpitaria)  uvasanus  (Conrad,  1855) 

Spondylus  carlosensis  Anderson,  1905 
Teredo?  sp. 

Vertebrata 

Isurus  cf.  I.  praecursor  (Leriche,  1906) 

Odontaspis  sp. 


turbidite-filled  incised  channels  were  covered  by  shallow- 
marine  facies  (Squires,  1981,  1983b)  (Fig.  2). 

The  coastal  alluvial-fan  facies,  equivalent  to  the  basal  con- 
glomerate depicted  in  Figure  1,  is  unfossiliferous.  The  zone 
of  interfingering  between  this  facies  and  the  shallow-marine 
facies  contains  fairly  abundant  mollusks.  Many  of  these  mol- 
lusks  are  the  same  species  that  occur  in  the  stratigraphically 
higher  shallow-marine  facies.  Some  of  the  mollusks,  how- 
ever, are  restricted  in  their  occurrence  in  the  Llajas  For- 
mation to  this  zone  of  interfingering.  These  mollusks  are 
Nerita  cf.  N.  ( Amphi nerita ) eorex,  V elates  perversus,  Turri- 
tella  meganosensis  protumescens,  Turritella  andersoni,  and 
Venericardia  ( Pacificor ) aragonia  joaquinensis.  Unidentifi- 
able ostreid  remains  also  characterize  this  zone  of  interfin- 
gering. At  CSUN  locality  542,  such  remains  form  an  ostreid 
coquinite.  Turritella  meganosensis  protumescens  typically 
occurs  as  robust,  thick-shelled  fragments  up  to  5.5  cm  in 
length.  This  taxon  belongs  to  the  T.  reversa  stock  of  Merriam 
(1941).  According  to  Saul  (1983),  the  presence  of  large  mem- 
bers of  this  stock,  as  well  as  the  presence  of  Venericardia 
(Pacificor)  aragonia  joaquinensis,  in  the  Llajas  Formation 


I’able  3.  Explanation  of  reference  abbreviations  for  Tables  4 and 
5. 


A1 

Abbott,  1968a 

A2 

Abbott,  1974 

A&D 

Abbott  and  Dance,  1982 

C 

Cossmann,  1901 

D 

Dance,  1976 

D&D 

Dell  and  Dance,  1963 

E 

Eisenberg,  1981 

G 

Grasse,  1968 

Ha 

Habe,  1968 

H&G 

Hertlein  and  Grant,  1972 

K 

Keen,  1971 

K&C 

Keen  and  Coan,  1974 

Ki 

Kira,  1965 

L 

Lindner,  1978 

Ma 

Marincovich,  1977 

Me 

McMillan,  1968 

Me 

Merriam,  1941 

Ml 

Morris,  1966 

M2 

Morris,  1975 

N 1 

Nordsieck,  1968 

N2 

Nordsieck,  1969 

O 

Olsson,  1961 

R 

Rios,  1970 

S 

Schenck,  1936 

T 

Tebble,  1976 

W&D 

Weaver  and  duPont,  1970 

indicate  nearshore,  shallow-water  conditions.  The  presence 
of  ostreid  coquinites  indicates  similar  conditions.  Nerita  oc- 
curs along  shorelines  today  (Tables  3 and  4),  and  Velates,  a 
closely  allied  but  extinct  genus,  probably  occurred  in  the  same 
type  of  environment. 

Most  of  the  fossils  in  the  formation  occur  in  the  shallow- 
marine  facies  proper,  which  can  be  divided  into  a transgres- 
sive phase  and  a regressive  phase.  The  transgressive  phase 
makes  up  most  of  the  lower  part  of  the  formation.  It  consists 
of  alternating  laminated  and  bioturbated  sandstone  with  scat- 
tered fossiliferous  beds  mostly  confined  to  channels.  The 
laminated  sandstone  represents  storm-influenced  stratifica- 
tion and  the  biogenic  reworking  is  inter-storm  activity 
(Squires,  1981).  These  deposits  probably  accumulated  where 
the  coastal  alluvial-fan  rivers  emptied  into  the  ocean.  The 
resulting  sediment  buildup  would  have  been  subjected  to 
reworking  by  waves  (Squires,  1 983b).  The  fossils  have  under- 
gone a small  distance  of  postmortem  transport  and  constitute 
indigenous  death  assemblages  (Squires,  1981).  Many  fossils 
show  preservation  of  delicate  features,  such  as  protoconchs, 
outer  lips,  and  ribs.  There  is  much  variability  in  the  taxo- 
nomic composition  of  the  fossils  in  these  channel-fill  depos- 
its. At  many  localities,  only  one  or  two  species  of  megafossils 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  5 


Table  4.  Recent  bathymetry  of  extant  molluscan  genera  of  the  Llajas  Formation. 


Genus  (subgenus) 

Depth  range 
(m) 

*MFR  depth 
(m) 

**References 

Scaphopoda 

Dentalium 

4-2320 

7-100 

A2,K,K&C,M2,R 

Gastropoda 

Ancilla 

6-87 

21-70 

A2,R 

Ancistrolepis 

40-200 

100 

K 

Architectonica 

0-134 

2-37 

A2,K 

Calyptraea 

0-70 

0-37 

A1,M1,M2,K 

Cerithiopsis 

0-823 

24-60 

A2.M1 

Cirsotrema 

33-274 

73-137 

A2,Ha,Ki 

Conus 

0-550 

9-55 

K,M  1 ,R 

Cymatium  (Septa) 

0-146 

24-60 

A2.M1.M2 

Cypraea 

0-140 

9-24 

A1,M1,M2,R 

Fusinus 

0-3616 

18-60 

A 1 ,A2,K,K&C,Ki,M  1 ,M2 

Fusiturricula 

56-65 

60 

R 

Galeodea 

7-330 

8 

Mc.Nl 

Lyria 

0-133 

2-24 

Ha,K,Ki,W&D 

Natica  (Naticarius) 

0-280 

32-60 

Ma,R 

Nerita 

0-9 

0-9 

K,M1,M2 

Never  it  a (Never it  a) 

1 1-2860 

658-1281 

M 

Olivella 

0-90 

0-27 

A1,A2,K,K&C,R 

Phalium  ( Semicassis ) 

18-100 

37-55 

Ha.Ki 

Polinices  (Euspira) 

0-4794 

15-200+ 

Ma,M  1 

Pseudoliva 

0-3 

0-3 

C,L 

Ranella 

55-915 

110 

D&D 

Scaphander 

20-5200 

50-160 

A1.A2.K 

Sinum 

0-170 

24-48 

K,K&C,Ma,R 

Terebra 

0-280 

4-18 

K,M  1 

Turritella 

4-185 

26-56 

K,K&C,Ki,Ml 

Xenophora 

0-200 

20-144 

A1  ,A2,Ha,K,Ki,M2,R 

Bivalvia 

Acanthocardia 

0-360 

5 

N2,T 

Acila  (Truncacila) 

7-1256 

20-100 

A1,A2 

Anomia 

0-550 

0-9 

A2,K,M1,M2 

Brachidontes 

0-31 

0-24 

K,M1,M2,R 

Callista  (Costacallista) 

30-214 

45-100 

A2,R 

Cardiomya 

4-2400 

60-190 

K,Ml,M2,Ha,R 

Corbula  (Caryocorbula) 

0-823 

10-27 

A2,K 

Gari 

0-150 

10-50 

A2,Ha,Ml 

Glycymeris 

0-110 

9-60 

A1,K,M1,M2 

Macoma 

0-1545 

0-45 

A1,K&C,M1,R 

Nemocardium 

9-640 

25-90 

A2,Ha,H&G,K,Ki,M  1 ,K.&C 

Nuculana  (Saccella) 

2-1097 

10-80 

A2,K 

Ostrea 

0-91 

0-24 

H&G,K,M1,M2 

Pinna 

0-200 

0-24 

A2,Ha,K,Ml,M2 

Pitar  (Lamelliconcha) 

0-110 

30-80 

K,0 

Solena 

Nearshore 

K 

Spondylus 

3-140 

15-50 

A1,K,M1,M2,R 

Teredo 

Bores  into  wood 

A2,M2 

* Most  frequently  reported;  **  see  Table  3 for  abbreviations. 


6 Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


are  present.  Turritella  andersoni  lawsoni  is  usually  one  of 
these  species.  At  other  localities,  such  as  CSUN  locality  371, 
as  many  as  45  species  of  megafossils  have  been  found. 

The  regressive  phase  of  the  shallow-marine  facies  makes 
up  the  uppermost  part  of  the  formation.  It  consists  of  bio- 
turbated  silty  sandstone  with  minor  occurrences  of  laminated 
sandstone  and  scattered  fossiliferous  units.  These  deposits 
were  in  slightly  deeper  waters  than  those  of  the  transgressive 
phase  and  were  less  affected  by  storm  reworking  (Squires, 
1983b).  Fossils  are  unabraded,  and  at  CSUN  locality  475 
articulated  Pinna  lewisi  specimens  were  found.  The  fossils 
in  the  regressive  phase  are  also  interpreted  to  be  indigenous 
death  assemblages. 

Where  the  sandstone  of  the  transgressive  and  regressive 
phases  of  the  shallow-marine  facies  grades  into  muddy  silt- 
stone  of  the  outer  shelf  and  slope  facies,  the  sandstone  is 
more  silty  and  more  bioturbated,  and  most  of  the  fossils  are 
confined  to  a few  beds  with  indistinct  contacts  due  to  bio- 
turbation  (Squires,  1981). 

One  such  bed,  informally  known  as  the  “Stewart  bed” 
(Squires,  1979,  1981,  1983b),  is  the  most  distinctive  fossil- 
iferous unit  in  the  shallow-marine  facies.  This  1-m-thick 
layer  occurs  in  the  uppermost  part  of  the  transgressive  phase 
of  the  shallow-marine  facies  in  the  northern  part  of  the  study 
area  and  can  be  traced  laterally  for  about  10  km  eastward 
from  the  type  section  of  the  formation.  Fossils  in  the  lower 
part  of  the  “Stewart  bed”  show  less  evidence  of  postmortem 
transport  than  those  elsewhere  in  the  shallow-marine  facies. 
Many  of  the  taxa  are  represented  by  nearly  complete  growth 
series  (with  only  the  early  juvenile  individuals  lacking).  Ar- 
ticulation of  valves  is  also  common.  There  is  a nearly  1:1 
ratio  of  opposite  valves  of  the  bivalves  Crassatella  uvasana 
and  Venericardia  (Pacificor)  hornii  calafia.  Some  specimens 
of  these  bivalves,  especially  Crassatella,  are  articulated  but 
are  not  in  growth  position.  Specimens  of  all  taxa  show  pres- 
ervation of  delicate  morphologic  features.  There  is  random 
orientation  of  the  faunal  remains.  Most  of  the  specimens 
occur  in  the  lower  50  cm  of  the  bed  and  seem  to  be  somewhat 
concentrated  in  indistinct  pods.  Based  on  the  above  features, 
as  well  as  the  lack  of  sedimentary  structures  indicative  of 
high  energy,  it  is  interpreted  that  a residual  (winnowed)  pa- 
leocommunity,  as  defined  by  Fagerstrom  (1964),  occurs  at 
many  exposures  of  the  lower  part  of  the  “Stewart  bed” 
(Squires,  1981). 

The  uniformity  of  the  lithology  throughout  the  “Stewart 
bed”  suggests  uniform  depositional  and  environmental  con- 
ditions. The  megafauna  is  also  fairly  uniform  throughout  the 
bed.  The  most  abundant  faunal  elements  are  carnivorous 
naticid  gastropods  ( Eocernina  and  Pachycrommium),  her- 
bivorous gastropods  (Turritella  and  Ectinochilus),  infaunal 
suspension  feeder  bivalves  ( Crassatella  and  Venericardia ), 
and  carnivorous  solitary  ahermatypic  scleractinian  corals 
( Trochocyathusl ).  This  Eocernina-Turrite/la-Crassatella- 
Trochocyathus ? paleocommunity  is  best  developed  at  and  in 
the  vicinity  of  CSUN  locality  374,  with  50  species  of  mega- 
fossils present.  The  shallow-marine  “Stewart  bed”  was  de- 
posited near  the  shelf/slope  break  based  on  calcareous  nan- 
nofossil  studies  (Filewicz  and  Hill,  1 983)  and  on  the  presence 


of  outer  shelf  and  slope  deposits  directly  above  it.  The  pres- 
ence of  abundant  large  specimens  of  Trochocyathusl  in  the 
“Stewart  bed,”  as  well  as  in  a few  thin  beds  a few  meters 
above  the  “Stewart  bed”  at  the  type  section,  is  consistent 
with  a shelf/slope  break  environment  for  this  part  of  the 
Llajas.  Solitary  ahermatypic  scleractinian  corals  tolerate  a 
wide  range  of  temperatures  and  live  at  all  depths,  attaining 
their  best  development  along  margins  of  the  continental 
shelves  (Wells,  1957;  Heckel,  1972). 

Megafossils  which  characterize  the  transgressive  phase  of 
the  shallow-marine  facies  are  Pseudophragmina  ( Proporo - 
cyclina)  c/arki,  Cylichnina  tantilla,  Ectinochilus  (Macilentos) 
macilentus,  Eocernina  hannibali,  Pachycrommium  clarki, 
Turritella  andersoni  lawsoni,  Turritella  buwa/dana,  Brachio- 
dontes  ( Brachidontes ) cowlitzensis,  and  Dent  ahum  ( Laevi - 
dentalium)  calafium.  The  only  taxon  that  characterizes  the 
regressive  phase  of  the  shallow-marine  facies  is  Macoma 
rosa. 

Megafossils  which  commonly  occur  in  both  the  transgres- 
sive and  regressive  phases  of  the  shallow-marine  facies  are 
Ca/yptrea  diegoana,  Phalium  ( Semicassis ) tuberculiformis, 
Turritella  uvasana  app/inae,  Corbula  (Caryocorbula)  dick- 
ersoni,  and  Glycvmeris  (Glycymeris)  rosecanyonensis. 

Many  of  the  molluscan  genera  of  the  shallow-marine  facies 
of  the  Llajas  are  extant  and  most  commonly  occur  today  in 
seas  less  than  80  m depth  (Tables  3 and  4).  The  discocyclinid 
foraminifers  lived  in  very  shallow  water  (below  tide  level  to 
perhaps  100  m)  (Vaughan,  1945).  Small  calcareous  benthic 
foraminifers  present  in  the  transgressive  and  regressive  phas- 
es of  the  shallow-marine  facies  of  the  formation  are  indicative 
of  inner  to  middle  neritic  conditions  (Schymiczek,  1983a, 
1983b). 

Megafossils  are  scarce  in  the  outer  shelf  and  slope  facies, 
but  foraminifers  indicative  of  bathyal  depths  are  common 
(Squires,  1981;  Schymiczek,  1983a,  1983b).  The  mollusks 
that  occur  in  this  facies  also  occur  in  the  shallow-marine 
facies.  Apparently,  many  of  the  outer  shelf  and  slope  facies 
mollusks  are  shallow-marine  species  that  were  transported 
into  the  deeper  water  facies.  This  is  especially  true  for  the 
mollusks  at  CSUN  locality  54 1 . This  locality  is  from  the  base 
of  a turbidite-filled  channel  in  the  outer  shelf  and  slope  facies. 
Some  of  the  fossils  occur  as  fragments. 

PALEOCLIMATE 

West  Coast  Eocene  megafauna  have  long  been  assigned  to 
tropical  or  subtropical  environments  (Arnold,  1909;  Dick- 
erson, 1917;  Smith,  1919;  Clark  and  Vokes,  1936;  Ber- 
thiaume,  1938;  Vokes,  1940;  Durham,  1950).  Among  the 
Eocene  molluscan  genera  that  Durham  (1950)  listed  as  par- 
ticularly characteristic  of  tropical  (20°C  or  warmer),  the  fol- 
lowing also  occur  in  the  Llajas  Formation:  the  gastropods 
Anci/la,  Architectonica,  Conus,  Cypraea,  Ectinochilus  [=Ri- 
mella ],  Eocernina,  Eocithara  [=Harpa],  Ficopsis,  Paraser- 
aphs  [=Terebellum\,  Pseudoliva,  Terebra,  Turritella,  Ve/ates, 
Xenophora,  the  bivalves  Corbula,  Pinna,  Pilar,  Spondylus, 
large  Venericardia,  Crassatella  [=Crassatellites\,  and  the 
cephalopod  Aturia. 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  7 


Table  5.  Recent,  marine  faunal  regions  (Ross,  1974:4)  of  extant  molluscan  genera  of  the  Llajas  Formation. 


Genus  (subgenus) 

Boreal 

Warm 

temperate 

Tropical 

*References 

Scaphopoda 

Dentalium 

X 

X 

M 

D,K,R 

Gastropoda 

A ncilla 

M 

A2,D,R 

A ncistrolepis 

M 

A&D,E,Ha,K 

Architectonica 

X 

M 

A2,K,M2 

Calyptraea 

X 

X 

M 

A1,D,K,M1,M2 

Cerithiopsis 

X 

X 

M 

A2,K,K&C,M  1 

Cirsotrema 

X 

M 

A2,Ha,Ki,K 

Conus 

X 

X 

M 

K,M  1 ,R 

Cymatium  (Septa) 

X 

M 

A2,D,E,K,M  1 ,M2 

Cypraea 

X 

M 

A1,K,L,M1,R 

Fusinus 

X 

X 

M 

Al,A2,D,K,Ki,M  1 ,M2 

Fusiturricula 

M 

A&D.R 

Galeodea 

M 

X 

D,E,Ha,Mc,N  1 

Lyria 

X 

M 

A2,Ha,K,Ki,W&D 

Natica  ( Naticarius ) 

X 

M 

Ma.R 

Nerita 

M 

K,L,M1,M2 

Neverita  (Neverita) 

M 

X 

M 

Olivella 

X 

X 

M 

A2,D,K,K&C,R 

Phalium  (Semicassis) 

X 

X 

M 

E,Ha,Ki,L 

Polmices  (Euspira) 

X 

X 

M 

Ma 

Pseudoliva 

M 

C,G,L 

Ranella 

X 

M 

D,E 

Scaphander 

X 

M 

X 

A 1 ,A2,D,K,M2 

Sinum 

X 

M 

K,K&C,Ma,R 

Terebra 

X 

M 

D,K,M  1 

Turrit  ella 

X 

M 

K,K&C,Ki,M  1 ,Me 

Xenophora 

X 

M 

A 1 ,A2,Ha,K,Ki,M2,R 

Bivalvia 

Acanthocardia 

X 

M 

D,E,L,N2,T 

Acila  (Truncacila) 

M 

M 

X 

A1,S 

Anomia 

X 

M 

X 

A2,D,K,L,M1,M2 

Brachidontes 

X 

X 

M 

D,K,M  1 ,M2,R 

Callista  (Costacallista) 

X 

M 

A2,L,R 

Cardiornya 

X 

X 

M 

Ha,K,Ml,M2,R 

Corbula  (Caryocorbula) 

X 

M 

A2,K 

Gari 

X 

X 

M 

A2,D,Ha,L,Ml 

Glycymeris 

X 

X 

M 

A1,K,M1,M2 

Macoma 

M 

M 

X 

A1,D,M1,R 

Nemocardium 

X 

X 

M 

A2,D,Ha,H&G,K,Ki,Ml 

Nuculana  (Saccella) 

X 

X 

M 

A2,K 

Ostrea 

X 

X 

M 

D,H&G,K,M  1 ,M2 

Pinna 

X 

X 

M 

A2,D,Ha,K,Ml,M2 

Pitar  (Lamelliconcha) 

M 

K.O 

Solena 

M 

K 

Spondylus 

X 

M 

D,K,M  1 ,M2 

Teredo 

X 

M 

X 

A2,M2 

x = Present,  M = Most  frequently  reported  occurrence.  * See  Table  3 for  abbreviations. 


8 Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Berthiaume  (1938)  and  Durham  (1950)  also  mentioned 
that  discocyclinid  foraminifers  are  indicative  of  tropical  or 
subtropical  environments.  All  the  recorded  species  of  Amer- 
ican Discocyclinidae  occur  in  tropical,  subtropical,  and  south 
temperate  latitudes  (Vaughan,  1945). 

More  recent  and  independent  evidence  supports  a tropical 
to  subtropical  environment  interpretation  for  West  Coast 
Eocene  faunas.  Mineralogical  analyses  of  kaolinite  and  quartz- 
rich,  buried-soil  profiles  in  northwest  Baja  California  and 
southwest  California  indicate  that  humid  tropical  climatic 
conditions  prevailed  during  Paleocene  to  mid-Eocene  time 
(Peterson  and  Abbott,  1979). 

Based  on  oxygen  isotope  compositions  of  Tertiary  planktic 
foraminifer  tests  throughout  the  world.  Savin,  Douglas,  and 
Stehli  (1975)  concluded  that  ocean  temperatures  remained 
warm  and  relatively  constant  from  Paleocene  through  middle 
Eocene  time.  Kennett  (1982)  has  similarly  stated  that  during 
the  Paleocene  and  Eocene,  world  climates  were  relatively 
warm  and  equable  with  low  pole-to-equator  temperature  gra- 
dients. He  further  stated  that  a late  Paleocene  warming  trend 
culminated  in  a period  of  peak  warming  during  the  early 
Eocene  to  early  middle  Eocene.  This  interval  of  time  coin- 
cides with  the  age  of  the  Llajas  Formation.  Such  equable 
conditions  ceased  in  earliest  Oligocene  time  with  the  ap- 
pearance of  a worldwide  cooling  trend.  This  cooling  was 
associated  with  the  development  of  circum-Antarctic  cir- 
culation (Kennett  et  ah,  1975;  Kennett,  1982;  Zinsmeister, 

1982) . 

Although  56  percent  of  the  molluscan  genera  collected  from 
the  shallow-marine  facies  of  the  Llajas  Formation  are  extant 
(Tables  3 and  5),  the  percentage  is  not  high  enough  to  permit 
accurate  generic  comparison  studies  with  modern  molluscan 
faunas  like  those  done  by  Addicott  (1970)  for  Oligocene, 
Miocene,  and  Pliocene  molluscan  faunas  of  the  West  Coast. 
Generic  comparison  studies  with  modem  molluscan  faunas 
are  also  difficult  to  do  with  early  Tertiary  faunas  because  of 
the  likelihood  of  significant  changes  in  environmental  tol- 
erance in  progressively  older  faunas  (Addicott,  1 970).  Never- 
theless, a general  comparison  with  modern  molluscan  faunas 
can  be  made.  Most  of  the  extant  molluscan  genera  of  the 
shallow-marine  facies  of  the  Llajas  most  commonly  occur 
today  in  tropical  seas  (Tables  3 and  5).  It  should  be  noted 
also  that  most  of  these  genera  can  occur  in  warm-temperate 
seas  as  well. 

The  taxa  Ancilla,  Pseudoliva,  Nerita,  Pitar  ( Lamel/icon - 
cha),  and  Solena  are  particularly  significant  as  they  are  con- 
fined to  tropical  waters  today.  In  addition,  several  of  the 
extinct  generic  and  subgeneric  taxa  in  the  Llajas  fauna,  in- 
cluding Ectinochilus,  Eocernina,  Eocithara  [=Harpa\,  Fi- 
copsis,  Gomphopages,  Laevityphis,  Lyriscapha  [=  Volutocris- 
tata ],  and  Exputens,  have  modern  analogues  that  are  also 
tropical  or  subtropical  in  distribution  (Clark  and  Vokes,  1936; 
Palmer,  1967;  Givens,  1974). 

The  presence  of  large  Turritella  ( i.e.,  T.  meganosensis  pro- 
tumescens)  in  the  Llajas  is  indicative  of  shallow  waters  that 
were  warmer  than  are  presently  found  at  this  latitude  (Saul, 

1983) . 

A small  percentage  of  the  Llajas  megafauna  does  have  a 


temperate  water  aspect  to  it.  Acanthocardia  and  Ranella  are 
most  commonly  found  in  temperate  seas  today.  Ancistrole- 
pis ? and  Neverita  ( Neverita ) are  strictly  cold-water  gastropods 
(Tables  3 and  5).  Acanthocardia  is  most  abundant  in  the 
“Stewart  bed,”  and  Ranella  and  Ancistrolepisl  were  found 
only  in  this  bed.  Neverita  (Neverita)  was  found  at  about  the 
same  horizon  as  the  “Stewart  bed.”  Most  likely,  their  pres- 
ence in  or  near  the  “Stewart  bed,”  which  was  deposited  near 
the  shelf/slope  break,  was  due  to  “tropical  submergence.” 
This  phenomenon  has  been  observed  in  modern  mollusks 
(Ekman,  1953).  Many  cold-water  mollusks  that  occur  in  shal- 
low water  at  higher  latitudes  have  been  found  to  inhabit  only 
deeper  water  of  similar  temperature  in  the  tropics. 

AGE 

The  Llajas  Formation  is  late  early  Eocene  through  early  mid- 
dle Eocene  in  age,  based  on  mollusks,  benthic  foraminifers, 
and  calcareous  nannofossils.  Assignment  to  European  Stan- 
dard Ages,  various  standard  plankton  zones,  and  West  Coast 
provincial  benthic  foraminifer  and  molluscan  stages  in  shown 
in  Figure  3. 

Prior  to  1936,  early  workers  using  mollusks  reported  the 
age  of  the  Llajas  Formation  as  Eocene  (Waring,  1914,  1917; 
Kew,  1919,  1924)  or  middle  Eocene  (Clark,  1921,  1926; 
Cushman  and  McMasters,  1936). 

Clark  and  Vokes  (1936)  informally  proposed  five  mollus- 
can provincial  Eocene  “Stages”;  namely  “Meganos,”  “Ca- 
pay,”  “Domengine,”  “Transition,”  and  “Tejon.”  They  rec- 
ognized two  faunal  zones  in  their  “Capay  Stage,”  and  they 
assigned  the  Llajas  Formation  to  the  upper  zone  of  the  “Ca- 
pay Stage”  through  the  “Transition  Stage.”  Givens  (1974) 
showed  that  their  upper  faunal  zone  of  the  “Capay”  should 
be  considered  part  of  the  “Domengine  Stage,”  and  he  re- 
stricted the  use  of  “Capay  Stage”  to  their  lower  faunal  zone 
of  the  “Capay  Stage.”  It  is  in  this  restricted  sense  that  “Capay 
Stage”  is  used  herein. 

Saul  (1983)  regarded  the  “Meganos  Stage”  as  late  Paleo- 
cene-early  Eocene,  the  restricted  “Capay  Stage”  of  Givens 
( 1 974)  as  early  Eocene,  the  “Domengine”  as  late  early  through 
early  middle  Eocene,  and  the  “Transition”  as  middle  Eocene. 
These  ages  are  used  for  this  present  report  (Fig.  3).  Weaver 
et  al.  ( 1 944)  regarded  the  “Tejon”  as  late  Eocene,  and  Givens 
and  Kennedy  (1979)  regarded  it  as  late  middle  Eocene  and/ 
or  late  Eocene.  Such  ages  are  used  for  this  present  report  (Fig. 
3). 

Merriam  and  Turner  (1937)  reported  that  in  the  Llajas 
Formation  the  “Capay  Stage”  is  represented  by  the  “basal 
conglomerate”  because  of  the  presence  of  Turritella  mega- 
nosensis protumescens,  a subspecies  regarded  by  them  as  an 
index  fossil  for  the  “Capay  Stage.”  Based  on  this  present 
study,  this  subspecies  occurs  only  where  the  basal  conglom- 
erate (coastal  alluvial-fan  facies)  interfingers  with  the  shal- 
low-marine facies. 

Vokes  (1939)  assigned  the  lower  part  of  the  Llajas  For- 
mation to  the  lower  “Capay  Stage”  based,  in  part,  on  the 
presence  of  Galeodea  sutterensis.  I was  unable  to  find  any  G. 
sutterensis  in  the  Llajas.  Vokes  did  not  figure  his  Galeodea 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  9 


species  from  the  Llajas  Formation,  nor  did  he  give  any  lo- 
cality information  or  catalog  numbers  of  the  specimen(s). 
Vokes  (1939)  assigned  the  middle  part  of  the  Llajas  For- 
mation to  the  upper  “Capay  Stage”  based  on  the  presence 
of  Galeodea  susanae,  which  can  be  documented  as  a Llajas 
species  (i.e.,  CSUN  locality  371).  The  “G.  susanae  Zone”  of 
Clark  and  Vokes  (1936)  was  originally  assigned  to  the  upper 
part  of  the  “Capay  Stage,”  but  it  was  shown  to  be  faunally 
indistinguishable  from  the  “Domengine  Stage”  by  Givens 
(1974)  and  was  reassigned  to  the  “Domengine  Stage.” 

Vokes  (1939)  also  used  the  presence  of  Fusiturricula  (Cre- 
naturricula)  crenatospira  in  the  “Lower  Llajas”  as  an  indi- 
cator of  the  “Capay  Stage.”  He  noted  the  presence  of  F.  (C.) 
crenatospira  domenginica  in  the  “Upper  Llajas”  and  consid- 
ered this  subspecies  confined  to  the  “Domengine  Stage.”  In 
this  present  study,  F.  (C.)  crenatospira  was  found  only  in  the 
“Stewart  bed”  (just  above  the  middle  of  the  formation)  where 
it  is  associated  with  F.  (C.)  crenatospira  domenginica.  F.  (C.) 
crenatospira  domenginica.  furthermore,  occurs  in  strata  be- 
low the  “Stewart  bed.” 

Merriam  (1941)  reported  the  presence  of  Turritella  an- 
dersoni  susanae  and  T.  uvasana  infera  from  the  “basal  con- 
glomerate” of  the  Llajas  Formation.  In  this  present  study, 
T.  andersoni  susanae  is  considered  equivalent  to  T.  ander- 
soni.  As  with  T.  meganosensis  protumescens,  these  taxa  occur 
in  the  Llajas  Formation  only  where  the  coastal  alluvial-fan 
facies  interfingers  with  the  shallow-marine  facies.  T.  ander- 
soni is  a “Capay  Stage”  index  fossil.  Although  it  has  not  been 
previously  reported  from  the  Llajas  Formation,  Weaver  et 
al.  (1944)  assigned  the  lower  part  of  the  Llajas  Formation  to 
their  “Capay”  Turritella  andersoni  Zone.  The  rest  of  the 
Llajas  Formation  was  assigned  to  the  “Domengine”  and  low- 
er “Transition”  “Stages”  by  Weaver  et  al.  (1944). 

Squires  (1981,  1983a,  1983b)  and  Saul  ( 1 983)  reported  the 
lowermost  mollusk-bearing  beds  of  the  Llajas  Formation  to 
be  late  early  Eocene  in  age  and  the  rest  of  the  fossil-bearing 
beds  as  early  middle  Eocene  in  age. 

Based  on  the  above  discussions,  it  is  concluded  that  the 
zone  of  interfingering  between  the  coastal  alluvial-fan  facies 
and  the  shallow-marine  facies  is  assignable  to  the  Turritella 
uvasana  infera  fauna  of  the  restricted  “Capay  Stage”  of  Giv- 
ens (1974)  which  is  equivalent  to  the  lower  of  the  two  faunal 
zones  of  the  “Capay  Stage”  of  Clark  and  Vokes  ( 1 936),  Vokes 
(1939),  and  Weaver  et  al.  (1944).  Due  to  the  presence  of 
many  “Domengine”  molluscan  species  associated  with  the 
“Capay”  species  where  the  coastal  alluvial-fan  facies  inter- 
fingers with  the  shallow-marine  facies,  it  is  probable  that 
most  of  the  “Capay  Stage”  is  not  represented  in  the  Llajas 
Formation.  The  part  that  is  present  is  probably  near  the 
“Capay-Domengine”  boundary.  Saul  (1983)  put  this  bound- 
ary near  the  P8-P9  boundary  of  the  “Standard”  planktic 
foraminiferal  zonation  scheme  as  used  by  Berggren,  Kent, 
and  Flynn  (in  press)  (Fig.  3).  Saul  (1983)  also  noted  that  most 
of  the  “Capay  Stage”  is  probably  missing  in  the  Simi  Valley 
area.  The  uppermost  “Capay  Stage”  part  of  the  Llajas  For- 
mation, therefore,  probably  corresponds  to  latest  early  Eocene 
time. 

Fossil-bearing  rocks  of  the  Llajas  Formation,  therefore. 


cannot  be  assigned  to  any  molluscan  stage  older  than  the 
Turritella  uvasana  infera  fauna  of  the  “Capay,”  as  used  in 
the  restricted  sense  of  Givens  (1974).  As  noted  by  Saul  (1983), 
no  mollusks  restricted  to  Clark  and  Vokes’  ( 1936)  older  “Me- 
ganos  Stage”  have  been  recovered  from  the  Llajas  Formation. 
Although  T.  uvasana  infera  and  T.  andersoni  susanae  have 
been  reported  from  the  upper  100  m of  the  Meganos-age 
Santa  Susana  Formation,  specimens  are  variants  slightly  dif- 
ferent from  those  at  the  type  localities  low  in  the  Llajas 
(Merriam,  1941;  Saul,  1983).  Filewicz  and  Hill  (1983)  re- 
ported a hiatus  of  approximately  4 million  years  between  the 
Llajas  Formation  and  the  underlying  Santa  Susana  Forma- 
tion. In  this  present  study,  this  hiatus  is  considered  to  be 
more  on  the  order  of  3 million  years  (Fig.  3). 

The  remaining  part  of  the  Llajas  Formation  is  associated 
with  megafossils  that  have  been  reported  (Vokes,  1939;  Mer- 
riam, 1941;  Verastegui,  1953;  Rehder,  1973;  Givens,  1974, 
1979;  Givens  and  Kennedy,  1979;  Saul,  1983)  elsewhere  on 
the  West  Coast  only  from  the  “Domengine  Stage.”  These 
taxa  are  Conus  caleocius,  Eocithara  mutica  californiensis, 
Fusinus  teglandae,  Fusiturricula  ( Crenaturricula ) crenato- 
spira domenginica.  Lyriscapha  lajollaensis , Molopophorus 
cretaceus,  Olequahia  domenginica,  Proximitral  cretacea, 
Pseudoliva  lineata,  Turritella  andersoni  lawsoni,  Turritella 
uvasana  applinae,  Xenophora  stocki,  Claibornites  diegoensis, 
Pitar  ( Lamelliconcha ) joaquinensis,  and  Venericardia  (Pa- 
cificor)  hornii  calafia. 

Additional  evidence  for  a “Domengine”  age  for  the  Llajas 
Formation  is  the  overlap  or  joint  occurrence  of  taxa  that 
have  been  reported  (Givens,  1974;  Givens  and  Kennedy, 
1979)  elsewhere  on  the  West  Coast  as  having  their  lowest 
stratigraphic  occurrence  in  the  “Domengine  Stage,”  includ- 
ing Ficopsis  cooperiana,  Laevityphis  (Laevityphis)  antiquus, 
Ranellina  pilsbryi,  Terebra  californica,  Glycymeris  ( Glycy - 
meris)  rosecanyonensis,  and  Glyptoactis  (Glyptoactis)  do- 
menginica, with  taxa  that  have  been  reported  (Givens,  1 974; 
Givens  and  Kennedy,  1979)  elsewhere  as  having  their  highest 
stratigraphic  occurrence  in  the  “Domengine  Stage,”  includ- 
ing Architectonica  (S  tel /axis)  cognata.  Ectinochilus  ( Maci - 
lentos)  macilentus,  Pseudoperissolax  blakei  praeblakei,  Ve- 
lates  perversus,  and  Nayadina  ( Exputens ) llajasensis. 

Based  on  calcareous  nannofossils  and  mollusks  in  the  Lla- 
jas Formation,  Saul  (1983)  reported  the  “Domengine  Stage” 
to  be  of  late  early  through  early  middle  Eocene  age. 

Strata  assignable  to  the  middle  Eocene  “Transition  Stage” 
were  not  recognized  in  the  Llajas  Formation  in  this  present 
study.  The  unique  association  of  molluscan  species  that  is 
utilized  in  recognizing  this  “stage”  (Givens,  1974;  Givens 
and  Kennedy,  1979)  was  not  found  in  the  Llajas  Formation. 

Workers  using  benthic  foraminifers  have  reported  the  age 
of  the  Llajas  Formation  as  middle  Eocene  (Cushman  and 
McMasters,  1 936)  or  early  to  middle  Eocene  (Laiming,  1 940a, 
1940b,  1943;  Mallory,  1959;  Schymiczek,  1983a,  1983b). 

Laiming  (1940a,  1940b,  1943)  assigned  the  Llajas  For- 
mation to  the  B-3  (upper  part),  B-2,  B-l,  and  B-1A  “Zones” 
of  his  tentative  benthic  foraminiferal  zonation  scheme.  He 
correlated  these  “zones”  to  the  upper  “Capay”  through  “Do- 
mengine” molluscan  “Stages”  of  Clark  and  Vokes  (1936). 


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Squires:  Simi  Valley  Eocene  Mollusks 


Mallory  (1959)  reported  that  the  lower  Llajas  corresponds 
to  his  Penutian  Stage  (lower  Eocene,  “Capay”)  and  that  the 
bulk  of  the  formation  corresponds  to  his  Ulatisian  Stage 
(middle  Eocene,  “Domengine  to  Transition”).  An  updated 
version  of  these  benthic  foraminiferal  stages,  now  known  to 
be  time-transgressive,  is  given  in  Poore  (1980).  Schymiczek 
(1983a,  1983b)  assigned  the  Llajas  Formation  to  the  Ulati- 
sian Stage.  Givens  and  Kennedy  (1979)  have  concluded  that 
the  “Domengine  Stage”  is  correlative  with  the  Ulatisian  (and 
probably  in  part  Penutian)  Stage. 

Using  the  calcareous  nannofossil  biostratigraphic  zones  of 
Okada  and  Bukry  (1980)  and  the  absolute  dates  assigned  to 
these  zones  by  Berggren,  Kent,  and  Flynn  (in  press),  Filewicz 
and  Hill  (1983)  reported  that  most  of  the  shallow-marine 
(transgressive)  facies  of  the  Llajas  Formation  is  of  latest  early 
Eocene  or  earliest  middle  Eocene  age  ( Discoaster  lodoensis- 
D.  sublodoensis  Zones,  CPI  1-CP12).  The  uppermost  30  m 
or  so  of  this  facies  (Discoasteroides  kuepperi  Subzone,  CP  1 2a) 
and  the  outer  shelf  and  slope  facies  are  of  middle  Eocene  age 
(Rhabosphaera  inflata  Subzone,  CP12b-CP13,  to  Nannotet- 
rina  quadrata  Zone,  CPI 3).  Rhabdosphaera  inflata  first  oc- 
curs 4 m above  the  “Stewart  bed.”  These  results  are  sum- 
marized in  Figure  3. 

According  to  Filewicz  and  Hill  (1983),  the  R.  inflata  Sub- 
zone plots  within  the  TE2. 1 global,  eustatic  sea-level  cycle 
of  Vail,  Mitchum,  and  Thompson  (1977),  and  the  sudden 
shift  to  upper  bathyal  waters  just  above  the  “Stewart  bed” 
may  be  in  response  to  both  a steady  increase  in  sea  level  and 
tectonism. 

CORRELATION 

Correlations  between  California  and  Oregon/Washington  can 
be  easily  accomplished  with  molluscan  assemblages  of  mid- 
dle and  early  late  Eocene  age  because  these  mollusks  occupied 
a single  faunal  province  (Armentrout,  1975). 

Twenty-two  (or  42  percent)  of  the  species  listed  by  Givens 
and  Kennedy  (1979)  for  the  “Domengine  Stage”  Mount  So- 
ledad,  Ardath  Shale,  and  Scripps  Formations  of  the  La  Jolla 
Group  near  San  Diego,  California  also  occur  in  the  Llajas 
Formation.  All  three  formations  yield  Turritella  andersoni 
lawsoni.  In  addition,  the  Ardath  Shale  yields  T.  uvasana 
applinae  and  has  planktic  foraminifers  correlative  with  P10/ 
1 1 Zones  and  calcareous  nannofossils  correlative  with  the 
CPI 2 Zone  (Givens  and  Kennedy,  1979). 

Most  of  the  megafauna  of  the  late  early  through  middle 
age  Eocene  Maniobra  Formation,  Orocopia  Mountains, 
southeastern  California  is  conspecific  with  that  of  the  Llajas 
Formation.  The  lower  part  of  the  formation  has  calcareous 
nannofossils  indicative  of  CP9  through  CPI  1 . Turritella  an- 
dersoni lawsoni  is  present  in  the  upper  part  of  the  formation 
(Crowell  and  Susuki,  1959;  Advocate,  1982). 

As  mentioned  earlier,  the  zone  of  interfingering  between 
the  coastal  alluvial-fan  facies  and  the  shallow-marine  facies 
of  the  Llajas  Formation  can  be  correlated  to  the  Turritella 
uvasana  infera  fauna  described  by  Givens  ( 1 974).  This  fauna 
is  from  the  lowermost  part  of  the  Juncal  Formation,  Pine 
Mountain  area,  California.  Fifteen  (or  40  percent)  of  the 


species  restricted  to  the  Turritella  uvasana  applinae  fauna 
described  by  Givens  (1974)  from  the  upper  half  of  the  Juncal 
Formation  in  the  same  area  occur  also  in  the  Llajas  For- 
mation above  the  zone  of  interfingering  between  the  coastal 
alluvial-fan  facies  and  the  shallow-marine  facies.  This  Juncal 
Formation  fauna  corresponds  to  the  upper  of  the  two  faunal 
zones  of  the  “Capay  Stage”  of  Clark  and  Vokes  ( 1 936)  through 
the  “Domengine  Stage”  (Givens,  1974,  fig.  6). 

Twenty-seven  (or  69  percent)  of  the  megafossil  species 
listed  by  Squires  (1977)  from  the  “Domengine-Transition” 
“Stages”  unnamed  stratigraphic  unit  E-,,  lower  Piru  Creek 
area,  Transverse  Ranges,  California,  also  occur  in  the  Llajas 
Formation. 

In  central  California,  the  megafaunas  of  both  the  Avenal 
Sandstone  (Stewart,  1946)  and  the  Domengine  Formation 
(Vokes,  1939)  are  similar  to  that  of  the  Llajas. 

In  southwestern  Oregon,  the  megafaunas  in  the  Umpqua 
and  Tyee  Formations,  as  used  by  Turner  (1938),  are  similar 
to  that  in  the  Llajas  Formation.  Turner  (1938:32)  assigned 
most  of  the  Umpqua  Formation  megafauna  to  the  “Capay 
Stage.”  He  locally  subdivided  the  Umpqua  into  lower  and 
upper  portions,  and  he  considered  the  upper  portion  of  his 
Glide  section  as  possibly  intermediate  between  the  “Capay” 
and  “Domengine”  “Stages.”  He  assigned  the  Tyee  mega- 
fauna to  the  “Domengine  Stage.” 

Turner’s  “lower  Umpqua”  now  corresponds  to  the  Rose- 
burg  Formation,  and  his  “upper  Umpqua”  includes  both  the 
Lookingglassand  Flournoy  Formations  (Baldwin,  1974).  The 
base  of  the  Lookingglass  Formation  is  considered  by  Baldwin 
to  coincide  with  the  division  between  Turner’s  “lower”  and 
“upper  Umpqua.”  Based  on  a study  of  planktic  foraminifers. 
Miles  (1981)  assigned  the  Roseburg  Formation  to  Zone  P7- 
8 of  the  “Standard”  zonation,  the  Lookingglass  Formation 
to  Zone  P7-8,  and  the  Flournoy  Formation  to  Zone  P10.  He 
assigned  the  Tyee  Formation  to  an  early  middle  Eocene  age. 

Miles  (1981),  in  his  study  of  the  southwestern  Oregon 
Eocene  strata,  placed  the  “Capay-Domengine”  boundary  near 
the  P9-10  boundary,  but  Saul  (1983)  has  pointed  out  that 
this  molluscan  stage  boundary  actually  should  lie  near  the 
P8-9  boundary.  Using  Miles’  (1981)  figures  3 and  7 and 
Saul’s  ( 1 983)  refinement  of  the  “Capay-Domengine”  bound- 
ary, the  Roseburg  and  Lookingglass  Formations  can  be  shown 
to  be  equivalent  to  the  “Capay  Stage”  and  the  Flournoy 
Formation  to  be  equivalent  to  the  “Domengine  Stage.” 

Miles  (1981,  fig.  7)  assigned  most  of  the  Llajas  Formation 
to  early  Eocene  time,  but  such  an  age  assignment  does  not 
conform  to  the  findings  of  this  present  report  or  to  the  findings 
of  Filewicz  and  Hill  (1983),  Schymiczek  (1983a,  1983b),  or 
Saul  (1983). 

BIOGEOGRAPHY 

The  widespread  aspect  of  West  Coast  warm-water  Eocene 
faunas  can  be  demonstrated  by  the  recognition  of  some  of 
Clark  and  Vokes’  (1936)  Eocene  “stages”  from  southern  Cal- 
ifornia (latitude  33°N)  to  as  far  north  as  the  Gulf  of  Alaska 
(latitude  60°N)  (Addicott,  1970).  About  30  percent  of  the 
Llajas  molluscan  species,  for  example,  ranged  from  San  Die- 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  11 


go  into  Oregon/Washington.  Two  other  species,  Acila  (Trun- 
cacila)  decisa  and  Nuculana  ( Saccella ) gabbii,  ranged  from 
southern  California  to  Kamchatka. 

Clark  and  Vokes  (1936)  used  the  presence  of  Velates  per- 
versus,  which  occurs  in  the  lower  part  of  the  Llajas,  as  evi- 
dence of  a seaway  connection  in  the  tropical  Central  Amer- 
ican region  between  the  Atlantic  Ocean  and  the  waters  of  the 
West  Coast  of  North  America.  Velates  is  predominantly 
Tethyan  in  its  distribution  and  has  been  found  in  Western 
Europe,  Africa,  Asia,  Burma,  West  Pacific,  Jamaica,  Florida, 
Panama,  and  southern  and  central  California  (Vokes,  1935; 
Clark  and  Vokes,  1936;  Palmer,  1967).  It  was  a shallow- 
water  mollusk  indicative  of  tropical  and  subtropical  climate, 
as  are  species  of  the  subgenus  Pacificor  of  the  bivalve  Ve- 
nericardia  (Palmer,  1967)  which  also  occur  in  the  Llajas. 

The  discocyclinid  foraminifer  Pseudophragmina  (Propo- 
rocyclina ) clarki  has  been  found  in  Florida,  Mexico,  and  Peru, 
as  well  as  along  the  West  Coast  (Cole  and  Applin,  1964; 
Blondeau  and  Brabb,  1983).  The  presence  of  this  species  in 
the  Llajas  Formation  is  additional  evidence  of  the  seaway 
connection  mentioned  above. 

Clark  and  Vokes  (1936)  also  cited  the  presence  of  Clavi- 
lithes  n.  sp.  A and  Cryptochorda  ca/ifornica  from  the  Llajas 
as  further  evidence  of  a Central  American  seaway  connection 
because  these  species  are  closely  related,  respectively,  to 
Clavilithes  scalaris  and  Cryptochorda  stromboides  from  the 
Paris  Basin  Eocene.  Other  Llajas  megafauna  species  used  as 
evidence  for  the  seaway  are  Clavilithes  tabulatus,  Lyria  an- 
dersoni,  and  Eocithara  mutica  californiensis,  which  are  closely 
related,  respectively,  to  Clavilithes  parisiensis,  Lyria  maga, 
and  Eocithara  mutica  of  the  Paris  Basin  Eocene  (Clark  and 
Vokes,  1936;  Vokes,  1937).  Some  other  Llajas  species  which 
have  been  noted  as  similar  to  ones  from  the  Paris  Basin 
Eocene  are  Callocardia  (Nitidavenus)  tejonensis  [=Isocardia] 
which  resembles  Cytherea  nitida  from  France  (Vokes,  1939) 
and  Benoistia  umpquaensis  which  closely  resembles  B.  brevi- 
cula  from  France  (Givens  and  Kennedy,  1976). 

Gardner  and  Bowles  (1934)  commented  upon  middle 
Eocene  gastropod  species  from  the  Chiapas  locality,  Isthmus 
of  Tehuantepec,  southern  Mexico  (Atlantic  side),  that  are 
closely  related  to  ones  from  the  Llajas  Formation.  These 
species  from  Chiapas  are  Cernina  ( Eocernina ) chiapasensis, 
Amaurellina  malinchae,  Amaurellina  cortezi,  and  Voluto- 
cristata  chiapasensis,  which  are  closely  related,  respectively, 
to  Eocernina  hannibali,  Tejonia  moragai,  Pachycrommium 
clarki,  and  Lyriscapha  lajollaensis  from  the  Llajas.  Gardner 
and  Bowles  (1934)  inferred  that  the  two  areas  were  once 
connected  by  a seaway. 

Givens  (1979),  in  a detailed  study  of  Lyrischapa,  conclud- 
ed that  this  genus,  which  is  present  in  the  shallow-marine 
(transgressive)  facies  of  the  Llajas,  probably  evolved  in  the 
Carribbean  region  during  Paleocene  or  early  Eocene  time.  It 
subsequently  spread  northward  during  the  middle  Eocene  or 
it  immigrated  from  the  Old  World  Tethyan  biogeographic 
province.  Entrance  into  the  Pacific  Coast  region  of  North 
America  was  by  way  of  a narrow  seaway  connection  across 
southern  Central  America  (Costa  Rica,  Panama)  or  north- 
western South  America  (Givens,  1979). 


According  to  Zinsmeister  (1983a),  the  first  indication  of 
the  influx  of  large  numbers  of  mollusks  from  the  Gulf  Coast 
and  the  Caribbean  into  western  North  America  was  in  the 
late  Paleocene,  and  this  influx  of  immigrants  continued  in 
the  early  Eocene.  “It  is  not  known  whether  these  immigrants 
came  in  distinct  pulses  associated  with  major  tectonic  events 
in  Central  America  or  represent  a continuous  long-term  west- 
ward migration  of  Caribbean  taxa  into  the  Pacific  as  a result 
of  a general  climatic  warming  during  the  Eocene”  (Zins- 
meister, 1983a:66). 

Most  of  the  megafauna  genera  of  the  Llajas  Formation 
were  already  present  on  the  West  Coast  by  Llajas  time.  In 
addition,  several  species  of  the  Llajas  megafauna  were  carry- 
overs from  earlier  times.  The  Llajas  taxa  Calyptraea  die- 
goana,  Pseudoperissolax  blakei  praeblakei,  Olivella  mathew- 
sonii.  Scaphander  ( Mirascapha ) costatus,  Surculites  mathew- 
sonii,  Acila  (Truncacila)  decisa,  Nemocardium  linteum,  and 
Nuculana  ( Saccella ) gabbii  have  been  reported  from  late  Pa- 
leocene “Martinez  Stage”  strata  on  the  West  Coast  (Nelson, 
1925;  Weaver,  1953;  Smith,  1 975;  Zinsmeister,  1974,  1983a). 
The  Llajas  taxa  Turbinolia  dickersoni,  Polinices  ( Euspira ) 
nuciformis,  and  Schizaster  diabloensis  have  been  reported 
from  late  Paleocene  strata  on  the  West  Coast  (Clark  and 
Woodford,  1927;  Quayle,  1932;  Marincovich,  1977).  The 
Llajas  taxa  Neverita  ( Neverita ) globosa,  Turritella  uvasana 
infera,  and  Brachidontes  ( Brachidontes ) cowlitzensis  have  been 
reported  from  late  Paleocene-early  Eocene  “Meganos  Stage” 
strata  on  the  West  Coast  (Givens,  1974;  Saul,  1983). 

Several  genera  immigrated  into  the  West  Coast  region  dur- 
ing Llajas  time.  Equatorial  paleocirculation  of  surface  waters 
was  still  largely  unrestricted  worldwide  during  Llajas  time 
and  until  at  least  45  million  years  ago  (middle  Eocene),  even 
though  the  Tethyan  Seaway  was  closing  up  due  to  compres- 
sive tectonic  forces  (Kennett,  1982).  As  mentioned  earlier, 
migration  from  points  east  of  the  West  Coast  was  accom- 
plished by  means  of  a narrow  seaway  across  southern  Central 
America  or  northwestern  South  America  (Woodring,  1966; 
Givens,  1977,  1979;  Zinsmeister,  1983a).  Molluscan  genera/ 
subgenera  present  in  the  Llajas  Formation  and  coeval  strata 
but  not  in  older  Tertiary  strata  of  the  West  Coast  will  be 
discussed  below.  Unless  otherwise  noted,  biogeographic  data 
for  these  genera  were  obtained  from  Wenz  (1938-1944), 
Palmer  and  Brann  (1965-1966),  Cox  et  al.  (1969),  Palmer 
(1974),  Marincovich  (1977),  and  Moore  (1983),  as  well  as 
from  the  “Systematics”  portion  of  this  present  report. 

Architectonica  ( Stellaxis ),  Paraseraphs,  and  Strepsidura 
possibly  arrived  during  the  early  Eocene  (uppermost  “Ca- 
pay”  part),  most  likely  from  the  Caribbean  province.  Do- 
menginella  apparently  originated  on  the  West  Coast  at  this 
time,  possibly  as  a northern  variant  of  Scobinella,  as  sug- 
gested by  Vokes  (1939). 

Benoistia  and  Megistostoma  arrived  during  the  late  early- 
early  middle  Eocene  (“Domengine”)  on  the  West  Coast  from 
the  Tethyan  province.  Eocithara  (see  Rehder,  1973),  Natica 
( Naticarius ),  Ranella  (see  Squires,  1983a),  and  possibly  Co- 
nus, Phalium  ( Semicassis ),  and  Marcia  (Mercimonia)  ar- 
rived during  this  time,  most  likely  from  the  Caribbean  region. 
Lyriscapha  (see  Givens,  1979)  may  have  arrived  from  either 


12  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


the  Caribbean  or  Tethyan  regions.  Cymatium  (Septa)  (see 
Squires,  1983a),  Galeodea  ( Ca/iagaleodea ),  Olequahia,  Te- 
jonia , Glyptoactis  (Glyptoactis),  and  possibly  Proxi mitral 
originated  on  the  West  Coast  at  this  time.  Ancistrolepisl  prob- 
ably arrived  during  the  early  middle  Eocene  from  the  Pacific 
Northwest. 

In  summary,  the  influx  of  mollusks  into  western  North 
America  that  Zinsmeister  ( 1 983a)  commented  on  continued, 
at  least,  into  the  early  middle  Eocene.  As  noted  by  Smith 
(1975),  the  megafauna  of  the  Llajas  Formation  is  of  mixed 
origins,  having  cosmopolitan,  Tethyan,  Caribbean,  and  North 
American  elements. 

SYSTEMATIC  MATERIALS  AND  METHODS 

From  1978  to  1983,  an  intensive  and  meticulous  search  of 
the  Llajas  Formation  was  undertaken  by  the  author  in  order 
to  amass  megafossil  specimens  that  would  represent  the  com- 
position and  the  stratigraphic  distribution  of  the  fauna  more 
accurately  than  the  cursory  collections  on  which  previous 
workers  based  their  reports.  About  5000  specimens  were 
obtained  from  132  localities,  permitting  the  addition  to  the 
known  fauna  of  many  rare  species  that  otherwise  would  not 
have  been  found.  Numerous  return  visits  were  made  to  sev- 
eral of  the  richest  localities.  Several  hundred  man-hours  were 
spent  quarrying  CSUN  locality  374  (“Stewart  bed”)  and 
CSUN  locality  371.  Because  specimens  with  preservation 
superior  to  those  previously  figured  were  obtained  and  be- 
cause many  of  the  holotypes  of  species  erected  for  Llajas 
Formation  specimens  are  from  localities  that  are  either  un- 
known or  only  vaguely  known,  most  of  the  holotypes  are  not 
refigured. 

Systematic  arrangement  of  the  generic  and  higher  taxo- 
nomic categories  follows  that  of  Cole  (1964)  for  the  large 
foraminifer;  Wells  (1956)  for  the  scleractinians;  Muir-Wood, 
Elliott,  and  Hatari  (1965)  for  the  brachiopod;  Howell  (1962) 
for  the  annelid  tube;  Ludbrook  (1960)  for  the  scaphopod; 
Cox  et  al.  (1969)  and  Vokes  (1980)  for  the  bivalves;  Miller 
(1947)  for  the  nautiloid;  Jeletzky  (1966)  for  the  sepiid; 
Glaessner  (1969)  for  the  brachyuran;  Fischer  (1966)  for  the 
echinoid;  and  Compagno  (1973)  for  the  sharks.  The  system- 
atic arrangement  of  Wenz  (1938-1944)  is  followed  generally 
for  the  gastropods. 

Most  of  the  figured  specimens  are  on  deposit  in  the  Natural 
History  Museum  of  Los  Angeles  County,  Invertebrate  Pa- 
leontology section.  Figured  specimens  borrowed  from  the 
paleontology  collections  of  the  University  of  California,  Los 
Angeles,  Department  of  Earth  and  Space  Sciences  are  on 
deposit  there.  Additional  unfigured  specimens  are  on  deposit 
in  the  Department  of  Geological  Sciences  Paleontology  col- 
lection, California  State  University,  Northridge. 

The  synonymies  are  selective.  Works  that  include  original 
figures  and/or  descriptions  are  listed.  References  that  add 
pertinent  and  documentable  biostratigraphic  information  are 
also  included.  More  complete  synonymies  of  many  of  the 
gastropods  and  bivalves  can  be  found  in  Stewart  (1927,  1 930). 

Primary  type  material,  molluscan  stage  range,  geographic 
distribution,  local  occurrence,  and  remarks  are  listed  for  all 


the  species.  Unless  otherwise  noted,  such  data  are  derived 
from  references  listed  in  the  synonymies  or  from  new  data 
obtained  in  the  course  of  this  present  study.  “Primary  type 
material”  refers  to  the  holotype,  paratype(s),  syntypes,  lec- 
totype,  paralectotype(s),  or  neotype  of  the  senior  subjective 
synonym  of  each  taxon.  In  the  case  of  homonyms,  the  junior 
homonym  “primary  type  material”  is  listed,  and  if  the  new 
name  “primary  type  material”  is  different  it  is  listed  also. 
The  molluscan  stages  are  for  the  West  Coast,  and  they  are 
from  Clark  and  Vokes  ( 1 936)  and  Weaver  et  al.  ( 1 944),  with 
refinements  made  by  Givens  (1974)  and  Saul  (1983).  The 
stages  are  provisional,  hence  the  names  are  placed  in  quo- 
tation marks.  The  relative  age  of  each  stage  and  correlation 
with  various  biostratigraphic  zones  are  shown  in  Figure  4. 
Any  taxa  stage  range  extensions  that  are  the  result  of  this 
present  study  of  the  Llajas  material  are  so  mentioned  under 
the  “Remarks”  for  each  species.  Locality  information  for  all 
the  localities  mentioned  in  this  report  is  given  in  the  “Lo- 
calities” section. 

Letter  abbreviations  used  for  catalog  and/or  locality  num- 
bers are: 

ANSP  = Academy  of  Natural  Sciences  of  Philadelphia 
CAS  = California  Academy  of  Sciences 
CIT  = California  Institute  of  Technology 
CSUN  = California  State  University,  Northridge 
MCZ  = Museum  of  Comparative  Zoology,  Harvard 
HS  = Herman  Schymiczek  (CSUN  master’s  thesis  field  sta- 
tion) 

LACMIP  = Los  Angels  County  Museum,  Invertebrate  Pa- 
leontology Section 

SU  = Stanford  University  (collections  now  housed  at  the 
California  Academy  of  Sciences) 

UCMP  = University  of  California  Museum  of  Paleontology 
(Berkeley) 

UCLA  = University  of  California,  Los  Angeles 
UCR  = University  of  California,  Riverside 
UO  = University  of  Oregon 

USGS  = United  States  Geological  Survey  (Washington,  D.C. 
register) 

USNM  = United  States  National  Museum  of  Natural  His- 
tory 

UW  = University  of  Washington 

SYSTEMATICS 
Phylum  Protista 
Subphylum  Sarcodina 
Class  Rhizopodea 
Subclass  Granuloreticulosia 
Order  Foraminiferida 
Suborder  Rotaliina 
Superfamily  Orbitoidacea 
Family  Discocyclinidae  Galloway,  1928 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  13 


Genus  Pseudophragmina  Douville,  1923 

Type  Species.  By  original  designation,  Orthophragmina 
flondana  Cushman,  1917. 

Pseudophragmina  ( Proporocyclina ) clarki 
(Cushman,  1920) 

Figures  5a-b 

Orbitolites  sp.  A Arnold,  1 9 1 0:pl.  3,  fig.  6. 

Orthophragmina  clarki  Cushman,  1920:41-42,  pi.  7,  figs. 
4-5. 

Discocyclina  clarki  (Cushman).  Schenck,  1929:221,  figs.  1- 
2,  5,  text  fig.  7.  Keenan,  1932:pl.  4,  figs.  1-2.  Vaughan, 
1936:255-256,  pi.  43,  figs.  1-2.  Berthiaume,  1938:496,  pi. 
61,  fig.  12.  Turner,  1938:7,  12,  21.  Stewart,  1946:table  1. 
Shepard,  Lankford,  and  Milow,  1957:columnar  section. 
Discocyclina  cloptoni  Vaughan.  Cushman  and  McMasters, 
1936:516,  pi.  77,  figs.  6-9.  [Misidentification  fide  Schy- 
miczek,  1983b:50.]  Vaughan,  1945:98-99,  pi.  42,  fig.  2. 
Pseudophragmina  ( Proporocyclina ) clarki  (Cushman). 
Vaughan,  1945:104,  106.  Cole,  1958:419-420,  pi.  52,  figs. 
3-11.  Crowell  and  Susuki,  1959:589.  Mallory,  1959:259. 
Cole  and  Applin,  1964:47,  pi.  9,  figs.  1-4.  Blondeau  and 
Brabb,  1983:47,  pi.  6,  fig.  17,  and  table  5. 
Pseudophragmina  clarki  (Cushman).  Schymiczek  and  Squires, 
1981:989.  Squires,  1983b:fig.  9i. 

Primary  Type  Material.  Holotype  and  paratype,  USNM 
collections,  upper  Lodo  Formation,  northeast  side  of  Do- 
mengine  Creek  near  the  comer  of  the  SW  Vi  of  section  29, 
T 18  S,  R 15  E,  Domengine  Ranch  quadrangle,  California. 
Molluscan  Stage  Range.  “Capay”  through  “Domengine.” 
Geographic  Distribution.  Florida,  Mexico,  Peru  through 
southwestern  Oregon. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  374,  457,  4691,  470b,  472,  473,  477,  479, 
487,  491,  504,  51  1,  522,  527a,  527b,  548. 

Remarks.  Specimens  are  essentially  confined  to  the  shal- 
low-marine (transgressive)  facies.  Most  occur  in  several  50- 
cm-thick  beds  approximately  100  m above  the  base  of  the 
formation  (e.g.,  localities  457,  4691,  487,  511,  527b).  In  such 
beds,  the  tests  are  concentrated  in  small  pods.  Turrit ella  an- 
dersoni  lawsoni  may  or  may  not  occur  with  the  tests,  but  if 
present  it  is  usually  the  only  associated  megafossil  or  the 
most  abundant  megafossil.  The  tests  of  P.  (P.)  clarki  are 
mostly  complete,  unabraded,  and  1.5  to  7 mm  in  diameter. 

The  stratigraphic  highest  occurrence  of  the  specimens  in 
the  Llajas  Formation  is  at  locality  473,  about  14  m above 
the  “Stewart  bed”  at  the  type  section.  The  bed  that  locality 
473  occurs  in  probably  represents  a shallow-marine  (trans- 
gressive) facies  bed  that  interfingers  with  the  outer  shelf  and 
slope  facies. 

Phylum  Coelenterata 
Subphylum  Cnidaria 
Class  Anthozoa 


Subclass  Zoantharia 
Order  Scleractinia 
Suborder  Caryophylliina 
Superfamily  Caryophylliicae 
Family  Caryophylliidae  Gray,  1847 
Subfamily  Caryophylliinae  Gray,  1847 

Genus  Trochocyathus 
Milne-Edwards  and  Haime,  1848 

Type  Species.  By  subsequent  designation  (Milne-Edwards 
and  Haime,  1850b),  Turbinolia  mitrata  Goldfuss,  1827. 

ITrochocyathus  striatus  (Gabb,  1864) 

Figure  5c 

Trochosmilia  striata  Gabb,  1864:207-208,  pi.  26,  fig.  195. 
Trochocyathus  striatus  (Gabb).  Vaughan,  1900:101-102,  pi. 

7,  fig.  24;  pi.  8,  figs.  1-3.  Dickerson,  1913:265.  Hanna, 

1927:269. 

Primary  Type  Material.  UCMP  holotype  12256,  Division 
B,  near  Mt.  Diablo,  California. 

Molluscan  Stage  Range.  “Capay”?;  Domengine. 

Geographic  Distribution.  Simi  Valley  through  central  Cal- 
ifornia. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  374,  438,  444,  445,  447,  451,  458,  459, 
471c,  472,  473,  477,  479,  480,  493,  494,  495,  496,  500,  510, 
525. 

Remarks.  ITrochocyathus  striatus  occurs  primarily  in  the 
“Stewart  bed.”  At  a few  localities  (e.g.,  locality  374)  there  is 
a partial  growth  series,  with  the  early  juveniles  missing.  Most 
specimens  in  the  Llajas  Formation  are  complete,  unattached, 
and  unabraded. 

Specimens  at  locality  473  occur  in  a bed  that  probably 
represents  a shallow-marine  facies  bed  that  interfingers  with 
the  outer  shelf  and  slope  facies. 

The  genus  and  species  of  the  most  abundant  coral  in  the 
Llajas  Formation  are  questioned  because  of  a lack  of  modem 
comparative  taxonomic  work  on  middle  Eocene  solitary 
scleractinian  ahermatypic  corals.  The  type  of  this  taxon  is 
largely  imbedded  in  matrix  and  probably  cannot  be  well 
enough  prepared  to  make  a valid  comparison  (Durham,  1981, 
pers.  commun.). 

Dickerson  (1913)  reported  T.  striatus  from  “Capay”-age 
strata  in  central  California,  but  he  gave  no  specific  locality 
information  nor  did  he  figure  the  species.  For  these  reasons, 
the  “Capay”  age  occurrence  is  uncertain. 

Subfamily  Turbinoliinae 
Milne-Edwards  and  Haime,  1848 

Genus  Turbinolia  Lamarck,  1816 

Type  Species.  By  subsequent  designation  (Milne-Edwards 
and  Haime,  1850b),  Turbinolia  sulcata  Lamarck,  1816. 


14  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Turbinolia  dickersoni  Nomland,  1916 

Figures  5d-e 

Turbinolia  dickersoni  Nomland,  1916:61,  pi.  3,  figs.  5-8. 

Palmer,  1923:306,  pi.  56,  figs.  2,  4.  Quayle,  1932:98-99, 

pi.  6,  figs.  1-5. 

Primary  Type  Material.  UCMP  holotype  12020  (missing 
since  1932),  Cerros  Shale  Member  of  the  Lodo  Formation, 
UCMP  locality  1817. 

Molluscan  Stage  Range.  Upper  Paleocene  through  “Do- 
mengine.” 

Geographic  Distribution.  Simi  Valley  through  central  Cal- 
ifornia. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  467. 

Remarks.  Only  four  specimens  were  found  in  the  Llajas 
Formation.  This  small  coral  is  distinguished  from  the  much 
larger  ITrochocyathus  striatus  by  the  presence  of  a stellate 
columella.  This  columella  does  not  project  noticeably  above 
the  corallum  wall. 

The  type  locality  of  this  species  is  in  the  lower  part  of  the 
Cerros  Shale  Member  (late  Paleocene  age)  or  the  Lodo  For- 
mation (Marincovich,  1977:252;  Moore,  1983:A88). 

Clark  (1926:114)  reported  this  species  from  “Domengine”- 
age  strata  in  central  California,  but  such  a report  is  uncertain 
because  of  his  lack  of  locality  data.  The  presence  of  this 
species  at  CSUN  locality  467  does  extend,  with  certainty,  its 
molluscan  stage  range  into  the  “Domengine  Stage.” 

Phylum  Brachiopoda 
Class  Articulata 
Order  Terebratulida 

Suborder,  Superfamily,  and  Family  UNKNOWN 
Genus  Eogryphus  Hertlein  and  Grant,  1 944 

Type  Species.  By  original  designation,  Eogryphus  tolmani 
Hertlein  and  Grant,  1944. 

Eogryphus  tolmani  Hertlein  and  Grant,  1944 

Figure  5f 

Eogryphus  tolmani  Hertlein  and  Grant,  1944:89-90,  pi.  5, 

figs.  1-3,  7;  pi.  18,  figs.  1,  9-11;  text  figure  22. 

Primary  Type  Material.  UCLA  holotype  6203,  CAS  para- 
types  7279-7282;  all  from  upper  Llajas  Formation,  upper 
Las  Llajas  Canyon,  3000  feet  north  and  1 500  feet  east  of  the 
southwest  corner  of  section  23,  T 3 N,  R 1 7 W (Hertlein  and 
Grant,  1944:89). 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  Simi  Valley,  California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  438,  468a,  515,  525. 

Remarks.  Many  specimens  were  found  at  and  near  the 
vicinity  of  locality  438.  Many  of  the  specimens  are  articu- 
lated. Plotted  on  the  topographic  map  available  in  1944,  the 
locality  of  the  type  specimens  probably  plots  about  1 500  feet 


(457  m)  east  of  CSUN  locality  5 1 5 in  an  area  in  which  slope 
wash  has  covered  the  exposures.  The  southwest  corner  of 
section  23  was  unmarked  and  had  to  be  projected.  The  lo- 
cality of  the  type  specimens,  however,  is  in  the  general  vi- 
cinity of  all  the  CSUN  localities  in  which  specimens  of  Eogry- 
phus tolmani  were  found. 

Smith  (1975:470)  reported  Eogryphus  cf.  E.  tolmani  from 
the  “Martinez  Stage”  (Paleocene)  part  of  the  Lodo  Forma- 
tion, central  California. 

Phylum  Annelida 
Class  Polychaetia 
Order  Sedentaria 

Family  Serpulidae  Lamarck,  1818 
Genus  Rotularia  Defrance,  1827 

Type  Species.  By  original  designation,  Serpula  spirulaea 
Lamarck,  1818. 

Rotularia  tejonense  (Arnold,  1910) 

Figure  5g 

Spiroglyphus ? tejonensis  Arnold,  1 9 1 0:5 1 , pi.  4,  fig.  18.  Dick- 
erson, 191 6:pl.  37,  figs.  5a-b.  Vokes,  1939:162-163,  pi. 

20,  figs.  20-22.  Stewart,  1946:pl.  11,  fig.  21. 

ITubulostium  tejonense  (Arnold).  Keen  and  Bentson,  1944: 

195. 

Rotularia  tejonense  { Arnold).  Nilsen,  1973:table  1.  Squires, 

1977:table  1 . 

Primary  Type  Material.  USNM  holotype  165658,  Avenal 
Formation,  USGS  locality  4617. 

Molluscan  Stage  Range.  “Capay”  through  “Transition.” 

Geographic  Distribution.  Simi  Valley  through  central  Cal- 
ifornia. 

Local  Occurrence.  Interfingering  coastal  alluvial-fan  facies 
and  shallow-marine  (transgressive)  facies:  CSUN  locality  703. 
Shallow-marine  (transgressive)  facies:  CSUN  localities  374, 
444,  445,  494,  521,  546. 

Remarks.  Most  specimens  were  found  in  the  “Stewart  bed” 
encrusting  other  shells,  most  notably  Ostrea. 

According  to  Clark  (1921:table  1),  R.  tejonense  [ =Spiro - 
glyphus  (?)  tejonensis]  occurs  in  “Meganos”  age  strata.  Such 
a report,  however,  cannot  be  substantiated  at  this  time  be- 
cause of  his  lack  of  locality  information  and  because  his  rock 
unit  names  are  not  clearly  defined. 

Phylum  Mollusca 
Class  Scaphopoda 
Family  Dentaliidae  Gray,  1834 
Genus  Dentalium  Linne,  1758 

Type  Species.  By  subsequent  designation  (Montfort,  1810), 
Dentalium  elephantinum  Linne,  1758. 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  15 


Dentalium  stentor  Anderson  and  Hanna,  1925 

Figure  5h 

Dentalium  stentor  Anderson  and  Hanna,  1925:145,  pi.  13, 

fig.  17. 

Dentalium  stentorl  Anderson  and  Hanna.  Squires,  1977 :ta- 

ble  1. 

Primary  Type  Material.  CAS  holotype  819,  Tejon  For- 
mation, CAS  locality  792. 

Molluscan  Stage  Range.  “Domengine”  through  “Tejon.” 

Geographic  Distribution.  Simi  Valley  through  southern  San 
Joaquin  Valley,  California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  373,  374,  444,  445,  447,  449,  451, 493, 
494,  500,  510. 

Remarks.  Abundant  large,  thick-shelled  specimens  were 
found  locally  in  the  “Stewart  bed”  (locality  374  and  vicinity). 
Although  some  specimens  are  nearly  complete  (up  to  75  mm 
in  height),  no  apices  were  found.  Due  to  the  absence  of  apices, 
the  specimens  could  not  be  assigned  with  certainty  to  any 
subgenus  of  Dentalium. 

Subgenus  Laevidenta/ium  Cossman,  1888 

Type  Species.  By  original  designation,  Dentalium  incertum 
Deshayes,  1825. 

Dentalium  ( Laevidenta/ium ) calafium 
Vokes,  1939 

Figure  5i 

Dentalium  (Laevidentalium)  calafium  Vokes,  1939:105,  pi. 

16,  figs.  30-31.  Squires,  1983b:fig.  9h. 

Primary  Type  Material.  UCMP  lectotype  15750,  Domen- 
gine  Formation,  UCMP  locality  A- 1027;  UCMP  paralecto- 
type  15751,  Domengine  Formation?,  UCMP  locality  A-1003. 

Molluscan  Stage  Range.  "Domengine.” 

Geographic  Distribution.  Simi  Valley  through  central  Cal- 
ifornia. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  37 1,373,  455,  466,  467,  484,  49 1 , 499,  500, 
507,  508,  510,  529b,  538,  539,  548. 

Remarks.  Specimens  occur  in  channel-fill  deposits  in  the 
Llajas  Formation.  They  are  small,  smooth,  and  fragmented 
specimens.  The  presence  of  the  apical  notch  on  the  convex 
side  of  the  shell  serves  to  distinguish  this  taxon. 

Vokes  (1939)  designated  two  syntypes  (UCMP  15750  and 
15751)  but  no  holotype  for  his  species.  In  accordance  with 
the  International  Code  of  Zoological  Nomenclature  (1964) 
article  74,  therefore,  UCMP  15750  is  herein  designated  as 
the  holotype  of  D.  (L.)  calafium. 

Class  Gastropoda 
Subclass  Prosobranchia 
Order  Archaeogastropoda 
Superfamily  Neritacea 


Family  Neritidae  Rafinesque,  1815 
Subfamily  Neritinae  Rafinesque,  1815 
Genus  Nerita  Linne,  1758 

Type  Species.  By  subsequent  designation  (Montfort,  1810), 
Nerita  peloronta  Linne,  1758. 

Nerita  cf.  N.  ( Amphinerita ) ear  ex 
Vokes,  1939 

Figure  6 a 

Local  Occurrence.  Interfingering  coastal  alluvial-fan  facies 
and  shallow-marine  (transgressive)  facies:  CSUN  localities 
700,  701. 

Remarks.  Specimens  were  found  only  in  the  “Capay  Stage” 
portion  of  the  Llajas  Formation.  A total  of  22  specimens  was 
found,  and  all  but  two  specimens  were  from  locality  700. 
Due  to  the  presence  of  well-cemented  matrix  in  the  aperture 
area  of  each  specimen,  the  labial  areas  could  not  be  studied. 
The  small  specimens  have  a very  low  spire,  large  body  whorl, 
and  a smooth  shell  surface.  Apparently,  there  is  no  umbilicus, 
and  the  outer  lip  is  not  dentate. 

The  specimens  most  closely  resemble  N.  (A.)  eorex  Vokes 
(1939:180-181,  pi.  22,  figs.  24,  26,  29)  from  the  Domengine 
Formation,  Reef  Ridge  area,  California.  Vokes  (1939)  re- 
garded this  species  as  the  only  smooth-shelled  Nerita  from 
the  West  American  Eocene. 

Genus  Velates  de  Montfort,  1810 

Type  Species.  By  original  designation,  Velates  conoideus 
Montfort,  1810  [=Nerita  perversa  Gmelin,  1791]. 

Velates  perversus  (Gmelin,  1791) 

Figures  6b-c 

Nerita  perversa  Gmelin,  1791:vol.  1,  pt.  6,  p.  3686. 

Velates  conoideus  Montfort,  1810:355. 

Velates  perversus  (Gmelin).  Vokes,  1935:382-383,  pi.  25, 

figs.  1-5;  pi.  26,  figs.  1-2.  Clark  and  Vokes,  1936:875,  pi. 

1,  figs.  7-8.  Givens,  1974:61,  pi.  5,  figs.  5-6,  13.  Givens 

and  Kennedy,  1979:83. 

Primary  Type  Material.  No  holotype  was  designated  by 
Gmelin  (1791).  Gmelin  did  not  have  a collection  but  worked 
mainly  from  the  literature,  according  to  Smith  (1970:459). 
Gmelin  (1791)  lists  the  Chemnitz  Cabinet  as  the  source  of 
his  Nerita  perversa,  and  if  the  Cabinet  is  still  extant,  the 
lectotype  could  be  designated. 

Molluscan  Stage  Range.  “Capay”  through  “Domengine.” 

Geographic  Distribution.  Western  Europe,  Africa,  Asia, 
Burma,  West  Pacific,  Jamaica,  southern  and  central  Califor- 
nia. 

Local  Occurrence.  Interfingering  coastal  alluvial-fan  facies 
and  shallow-marine  (transgressive)  facies:  CSUN  localities 
700,  701,  702. 

Remarks.  Specimens  were  found  only  in  the  “Capay  Stage” 
portion  of  the  Llajas  Formation.  Except  at  locality  701,  only 
fragments  were  found.  A very  large  specimen  (diameter  70 


16  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Figure  5.  Llajas  Formation  large  benthic  forammifer,  solitary  corals,  brachiopod,  calcareous  worm  tube,  and  scaphopods.  a and  b.  Pseudo- 
phragmina  ( Proporocyclina ) clarki  (Cushman,  1920).  a,  test  exterior,  LACMIP  hypotype  6500,  CSUN  loc.  511,  x 8.  b,  equatorial  section, 
LACMIP  hypotype  6501,  HS  loc.  HS-A-1 1(4),  x 25.  c.  ITrochocyathus  striatus  (Gabb,  1864),  lateral  view,  LACMIP  hypotype  6502,  CSUN 
loc.  374,  x 1.  d and  e.  Turbinolia  dickersoni  Nomland,  1916,  LACMIP  hypotype  6503,  CSUN  loc.  467.  d,  dorsal  view,  x 7.  e,  lateral  view, 
x 5.  f.  Eogryphus  tolmani  Hertlein  and  Grant,  1944,  dorsal  view,  LACMIP  hypotype  6504,  CSUN  loc.  438,  x 0.75.  g.  Rotularia  tejonense 
(Arnold,  1910),  LACMIP  hypotype  6505,  CSUN  loc.  444,  x 1.5.  h.  Dentalium  stentor  Anderson  and  Hanna,  1925,  partial  specimen,  side 
view,  LACMIP  hypotype  6506,  CSUN  loc.  444,  x 1 . i.  Dentalium  ( Laevidentalium)  calafium  Vokes,  1939,  partial  specimen,  apical  notch 
view,  LACMIP  hypotype  6507,  CSUN  loc.  373,  x 4.5. 


cm)  was  found  at  locality  701.  The  best  specimens  from  the 
Llajas  are  in  the  UCMP  collection  and  are  from  a locality 
equivalent  to  CSUN  locality  702  (i.e.,  UCMP  locality  7193). 
UCMP  specimen  37435  (Figs.  6b-c)  is  from  this  locality. 
Vokes  (1935)  figured  two  hypotypes  from  this  same  locality. 

The  most  characteristic  feature  of  this  species  is  the  pres- 
ence of  seven  or  eight  small  teeth  on  the  inner  lip. 

Order  Mesogastropoda 

Superfamily  Cerithiacea 

Family  Turritellidae  Woodward,  1851 

Genus  Turritella  Lamarck,  1799 

Type  Species.  By  monotypy,  Turbo  terebra  Linne,  1758. 

Turritella  meganosensis  protumescens 

Merriam  and  Turner,  1937 

Figure  6d 

Turritella  meganosensis  n.  subsp.  Clark,  1929:pl.  10,  figs. 
1-2. 


Turritella  meganosensis  protumescens  Merriam  and  Turner, 

1937:104,  pi.  6,  figs.  8-10.  Turner,  1938:85,  pi.  22,  fig.  15. 

Merriam,  1941:75-76,  pi.  8,  figs.  1-2,  5,  6,  8.  Weaver, 

1943:369-370,  pi.  74,  figs.  14,  18.  Saul,  1983:pl.  2,  fig.  1. 

Primary  Type  Material.  UCMP  holotype  15353,  “basal 
conglomerate”  of  the  Llajas  Formation,  UCMP  locality  7195. 

Molitiscam  Stage  Range.  “Capay.” 

Geographic  Distribution.  Simi  Valley,  California  through 
southwestern  Oregon. 

Local  Occurrence.  Interfingering  coastal  alluvial-fan  facies 
and  shallow-marine  (transgressive)  facies:  CSUN  localities 
452,  454,  469a,  469b,  469c,  501,  542,  545. 

Remarks.  According  to  Merriam  and  Turner  (1937:104), 
this  taxon  occurs  in  the  basal  conglomerate  of  the  Llajas 
Formation.  This  is  misleading,  as  specimens  do  not  occur  in 
the  basal  conglomerate  proper  (equivalent  to  the  coastal  al- 
luvial-fan facies).  They  actually  are  confined  to  beds  that 
represent  shallow-marine  facies  interfingering  with  the  coast- 
al alluvial-fan  facies.  At  a few  of  the  localities,  the  thick- 
shelled,  robust  T.  meganosensis  protumescens  exhibits  growth 
series. 


Contributions  in  Science,  Number  350 


Squires:  Simi  Vaiiey  Eocene  Mollusks  17 


Turritella  andersoni  Dickerson,  1916 

Figure  6e 

Turritella  andersoni  Dickerson,  1916:501-502,  pi.  42,  figs. 
9a-b.  Turner,  1938:83,  pi.  22,  figs.  4-6.  Merriam,  1941: 
76-77,  pi.  9,  figs.  1-2;  pi.  10,  figs.  1,  3-5,  8;  pi.  12,  figs. 
1-3.  Givens,  1974:62,  pi.  5,  figs.  7-10. 

Turritella  andersoni  susanae  Merriam,  1941:79,  pi.  11,  fig. 
6.  Saul,  1983:pl.  2,  fig.  5. 

Primary  Type  Material.  UCMP  holotype  12131,  Domen- 
gine  Formation,  UCMP  locality  1817. 

Molluscan  Stage  Range.  Upper  “Meganos”?,  “Capay.” 
Geographic  Distribution.  Simi  Valley,  California  through 
southwestern  Oregon. 

Local  Occurrence.  Upper  part  of  Santa  Susana  Formation?; 
interfingering  coastal  alluvial-fan  facies  and  shallow-marine 
(transgressive)  facies  of  the  Llajas  Formation:  CSUN  local- 
ities 469d,  700. 

Remarks.  Only  a single  specimen  was  found  at  locality 
469d.  This  specimen  (Fig.  6e)  consists  of  only  2Vi  whorls, 
but  the  characteristic  noded  three  primary  spiral  ribs,  the 
median  one  situated  closer  to  the  anterior  than  the  posterior 
primary,  are  present. 

Although  the  holotype  (UCMP  15295,  UCMP  locality 
A-993)  of  T.  a.  susanae  is  from  strata  equivalent  to  the 
interfingering  coastal  alluvial-fan  facies  and  shallow-marine 
(transgressive)  facies,  specimens  have  been  reported  by  Mer- 
riam (1941)  from  the  upper  part  of  the  underlying  Santa 
Susana  Formation.  Saul  (1983)  identified  these  specimens 
with  T.  andersoni  n.  sp.  because  none  has  the  sculpture  of 
the  Llajas  specimens.  Saul  (1983),  however,  did  note  that 
some  of  the  Santa  Susana  specimens  are  very  close  to  T. 
andersoni  Dickerson,  especially  from  localities  near  the  Lla- 
jas-Santa  Susana  contact. 

Turritella  andersoni  lawsoni  Dickerson,  1916 

Figures  6f-g 

Turritella  lawsoni  Dickerson,  1916:502,  pi.  42,  figs.  lOa-b. 
Hanna,  1927:308,  pi.  49,  fig.  5.  Hanna  and  Hertlein,  1943: 
fig.  62-15.  Stewart,  1946:pl.  11,  figs.  27-29. 

Turritella  andersoni  Dickerson,  Waring,  1917 :pl.  15,  fig.  18. 
Turritella  andersoni  lawsoni  Dickerson.  Vokes,  1939:161. 
Merriam,  1941:77-78,  pi.  9,  figs.  3-8;  pi.  12,  fig.  4.  Crowell 
and  Susuki,  1959:pl.  2,  fig.  12.  Givens,  1974:62-63,  pi.  5, 
figs.  11-12,  14.  Givens  and  Kennedy,  1979:82-83,  table 
1.  Saul,  1983:pl.  2,  figs.  10-1  1.  Squires,  1983b:fig.  9e. 
Turritella  andersoni  lawsoni  secondaria  Merriam,  1941:78- 
79,  pi.  9,  fig.  9.  Crowell  and  Susuki,  1959:pl.  2,  fig.  11. 
Turritella  andersoni  lawsoni  forma  secondaria  Merriam.  Saul, 
1983:pl.  2,  fig.  12. 

Primary  Type  Material.  UCMP  holotype  12128,  Domen- 
gine  Formation,  UCMP  locality  2295. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  San  Diego  through  central  Cal- 
ifornia. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 


CSUN  localities  374,  439,  440,  441, 442,  443,  444,  445,  447, 
449,  453,  455,  457,  468b,  469j,  470c,  471a,  471b,  471e,  481, 
485,  487,  490,  493,  494,  510,  516,  522,  523,  527a,  529a, 
529b,  537,  539,  548,  704. 

Remarks.  Turritella  andersoni  lawsoni  is  one  of  the  most 
abundant  megafaunal  components  in  the  Llajas  Formation. 
At  many  localities  (e.g.,  442  and  471a),  it  is  the  only  mega- 
fossil present.  T.  andersoni  lawsoni  is  restricted  in  the  Llajas 
Formation  to  the  shallow-marine  (transgressive)  facies.  Its 
highest  stratigraphic  occurrence  is  the  “Stewart  bed,”  where 
it  is  abundant.  Some  specimens  show  faint  spiral  ribs,  where- 
as others  do  not.  Some  specimens,  especially  at  locality  374, 
have  the  spiral  ribs  on  one  side  but  not  on  the  other  (weath- 
ered) side. 

Some  specimens  of  T.  a.  lawsoni  from  localities  374  and 
548  are  characterized  by  the  presence  of  a well-developed 
anterior  primary  rib  and  represent  the  form  described  by 
Merriam  (1941:78-79,  pi.  9,  fig.  9)  as  variety  secondaria. 
These  Llajas  specimens  of  T.  a.  lawsoni  which  show  the  well- 
developed  anterior  primary  rib  are  probably  just  ecotypes. 

The  primary  type  material  of  secondaria  is  UCMP  33998 
from  UCMP  locality  7004  = CSUN  locality  374. 

Turritella  buwaldana  Dickerson,  1916 

Figure  6h 

Turritella  buwaldana  Dickerson,  1916:500-501,  pi.  42,  figs. 

7a-b.  Hanna,  1927:307,  pi.  49,  figs.  7-8,  12.  Merriam, 

1941:86-87,  pi.  21,  figs.  3-9;  pi.  22,  figs.  1-14.  Stewart, 

1946:pl.  11,  fig.  24.  Givens,  1974:63,  pi.  5,  fig.  15.  Saul, 

1983:pl.  2,  figs.  13-15.  Squires,  1983b:fig.  9f. 

Turritella  kewi  Dickerson,  1916:501,  pi.  42,  fig.  8. 

Primary  Type  Material.  UCMP  holotype  12130,  Domen- 
gine Formation,  UCMP  locality  672. 

Molluscan  Stage  Range.  Upper  “Meganos”?,  “Capay”?, 
“Domengine”  through  “Tejon.” 

Geographic  Distribution.  San  Diego,  California  through 
southwestern  Oregon. 

Local  Occurrence.  Upper  part  of  the  Santa  Susana  For- 
mation?; shallow-marine  (transgressive)  facies  of  the  Llajas 
Formation:  CSUN  localities  371,  372,  373,  455,  467,  469e, 
469h,  469j,  484,  486,  499,  500,  505,  514,  538,  539,  540, 
548,  705. 

Remarks.  T.  buwaldana  in  the  Llajas  Formation  is  char- 
acterized by  its  small  size  and  presence  of  three  primary  spiral 
ribs  and  two  posterior  secondaries. 

T.  buwaldana  has  been  reported  from  the  upper  part  of 
the  Santa  Susana  Formation  by  Merriam  (1941),  but  he  noted 
that  these  forms  are  probably  subspecifically  distinct  from 
those  in  the  Llajas  Formation. 

Saul  (1983)  reported  T.  b.  crooki  Merriam  and  Turner 
from  the  “Capay”  portion  of  the  Llajas  Formation.  Although 
none  was  found  in  the  present  study,  it  is  possible  that  T.  b. 
crooki  and  T.  buwaldana  are  the  same.  More  taxonomic  data 
are  needed  to  resolve  the  question  (Saul,  1983,  pers.  com- 
mun.). 


18  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Turritella  uvasana  infera  Merriam,  1941 

Figure  6i 

Turritella  uvasana  infera  Merriam,  1941:90,  pi.  40,  figs.  2- 
4.  Givens,  1974:65-66,  pi.  6,  figs.  5-7.  Saul,  1983:pl.  2, 
fig.  4. 

Primary  Type  Material.  UCMP  holotype  33993,  “basal 
conglomerate”  of  the  Llajas  Formation,  UCMP  locality 
A-994  = CSUN  locality  452;  UCMP  paratypes  1 5439  and 
1 5443,  upper  part  of  the  Santa  Susana  Formation,  UCMP 
locality  7000. 

Molluscan  Stage  Range.  Upper  “Meganos”  through  “Ca- 
pay.” 

Geographic  Distribution.  Simi  Valley  and  Pine  Mountain 
area,  southern  California. 

Local  Occurrence.  Upper  part  of  the  Santa  Susana  For- 
mation; interfingering  coastal  alluvial-fan  facies  and  shallow- 
marine  (transgressive)  facies  of  the  Llajas  Formation:  CSUN 
localities  452,  705. 

Remarks.  According  to  Merriam  (1941:90),  this  taxon  oc- 
curs in  the  basal  conglomerate  of  the  Llajas  Formation.  This 
is  misleading,  as  specimens  do  not  occur  in  the  basal  con- 
glomerate proper  (equivalent  to  the  coastal  alluvial-fan  fa- 
cies). They  actually  are  confined  to  beds  that  represent  shal- 
low-marine facies  interfingering  with  the  coastal  alluvial-fan 
facies. 

T.  u.  infera  has  been  reported  from  the  upper  part  of  the 
Santa  Susana  Formation,  but  the  specimens  are  slightly  dif- 
ferent variants  that  have  more  rounded  whorl  profiles  and 
heavier  ribbing  than  those  from  the  type  locality  in  the  Llajas 
Formation  (Merriam,  1941;  Saul,  1983). 

Adult  whorls  of  the  Llajas  specimens  have  six  to  seven 
primary  spiral  ribs  with  the  posteriormost  two  more  closely 
spaced  and  somewhat  weaker  than  the  other  spiral  ribs.  Sec- 
ondary ribs  are  generally  absent. 

The  illustrated  specimen  (Fig.  6i)  is  the  same  one  used  by 
Saul  (1983:pl.  2,  fig.  4). 

Turritella  uvasana  applinae  Hanna,  1927 

Figure  6j 

Turritella  applini  Hanna,  1927:307,  pi.  49,  figs.  1,  4.  Clark, 
1929:pl.  10,  figs.  8,  18. 

Turritella  uvasana  applini  Hanna.  Merriam,  1941:93-94,  pi. 
16,  figs.  5-6;  pi.  18,  fig.  2. 

Turritella  uvasana  etheringtoni  Merriam,  1941:94,  pi.  15, 
figs.  12-15.  Squires,  1977:table  1. 

Turritella  uvasana  applinae  Hanna.  Givens,  1974:66,  pi.  6, 
figs.  3-4;  pi.  7,  fig.  19.  Givens  and  Kennedy,  1979:82-83, 
table  1.  Saul,  1983:pl.  2,  fig.  18.  Squires,  1983b:fig.  9g. 

Primary  Type  Material.  UCMP  holotype  30971,  La  Jolla 
Formation,  UCMP  locality  3993;  UCMP  paratype  33894, 
La  Jolla  Formation,  UCMP  locality  3990. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  San  Diego  through  Pine  Moun- 
tain area,  southern  California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 


CSUN  localities  371,  374,  440,  444,  449,  469e,  469i,  47 Id, 
471e,  481,  484,  486,  491,  500,  507,  529a,  539,  548,  704. 
Outer  shelf  and  slope  facies:  CSUN  locality  483.  Shallow- 
marine  (regressive)  facies:  CSUN  localities  475,  489,  512b, 
51 2d. 

Remarks.  Llajas  specimens  of  T.  u.  infera  superficially  re- 
semble T.  u.  applinae,  but  the  latter  has  a more  convex  whorl 
profile,  many  more  primary  spiral  ribs,  and  the  presence  of 
numerous  well-developed  secondary  ribs. 

The  primary  type  material  for  T.  u.  etheringtoni  is  from 
the  Llajas  Formation:  UCMP  holotype  33875,  UCMP  lo- 
cality 7003;  UCMP  paratypes  33876  and  33877,  UCMP  lo- 
cality 7004  = CSUN  locality  374.  Specimens  of  T.  u.  appli- 
nae are  especially  abundant  at  locality  374. 

Family  Architectonicidae  Gray,  1850 
Genus  Architectonica  Roding,  1798 

Type  Species.  By  subsequent  designation  (Gray,  1847), 
Trochus  perspectivus  Linne,  1758. 

Subgenus  Architectonica  s.s. 

Architectonica  {Architectonica)  llajasensis 
Sutherland,  1966 

Figure  6k 

Architectonica  llajasensis  Sutherland,  1966:1-4,  figs.  1-2. 

Primary  Type  Material.  LACMIP  holotype  1140,  Llajas 
Formation,  LACMIP  locality  461-B. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  Simi  Valley,  California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  371. 

Remarks.  Only  three  specimens  were  found,  and  these  are 
from  the  locality  in  Devil  Canyon.  Judging  from  the  mega- 
fauna associated  with  Sutherland’s  (1966)  holotype  speci- 
men, the  type  locality  of  A.  llajasensis  is  probably  near  CSUN 
locality  444  in  the  “Stewart  bed.” 

This  species  is  placed  in  Architectonica  s.s.  because  the 
umbilicus  is  bordered  by  two  serrated  ridges  (see  Sutherland, 

1 966),  a characteristic  of  typical  Architectonica  (Stewart,  1 927: 
343). 

Subgenus  Stellaxis  Dali,  1892 

Type  Species.  By  original  designation,  Solarium  alveatum 
Conrad,  1833. 

Architectonica  ( Stellaxis ) cognat  a 

Gabb,  1864 

Figure  61 

Architectonica  cognata  Gabb,  1864:1 17,  pi.  20,  figs.  72,  72a, 
72c,  not  d and  e as  stated  [not  72b  = A.  alveata  (Conrad) 
fide  Stewart,  1927:344], 

Stellaxis  cognata  (Conrad).  Waring,  1917:98. 

Architectonica  cognata  Gabb.  Givens  and  Kennedy,  1979: 
83,  table  1 . 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  19 


Architectonica  (Stellaxis)  cognata  Gabb.  Stewart,  1927:343- 
344,  pi.  28,  figs.  7-8.  Turner,  1938:90,  pi.  18,  fig.  17.Vokes, 
1939:163-164.  Stewart,  1946:pl.  11,  fig.  4.  Weaver,  1943: 
363-364,  pi.  73,  fig.  20;  pi.  103,  fig.  19.  Givens,  1974:68, 
pi.  7,  figs.  1-3. 

Primary  Type  Material.  ANSP  lectotype  4223,  Tejon  For- 
mation s.l.,  1 1 km  south  of  Martinez,  California. 

Molluscan  Stage  Range.  Uppermost  “Capay”  through 
“Domengine.” 

Geographic  Distribution.  San  Diego,  California  through 
southwestern  Oregon. 

Local  Occurrence.  Interfingering  coastal  alluvial-fan  facies 
and  shallow-marine  (transgressive)  facies:  CSUN  localities 
452,  702,  703.  Shallow-marine  (transgressive)  facies:  CSUN 
localities  371,  372,  373,  374,  444,  447,  455,  486,  490,  491, 
538,  704. 

Remarks.  This  species  is  characterized  mostly  by  a single 
keel  on  the  periphery  of  the  whorls  (Givens,  1974:68).  The 
Llajas  specimens  have  this  prominent  keel,  as  well  as  a less 
prominent  sutural  cord,  thereby  producing  an  apparent  dou- 
ble-keel look.  The  sutural  cord  is  exposed  due  to  the  removal 
of  the  shell  material  in  the  suture  region. 

A previously  assigned  “Capay”  age  (Turner,  1938;  Givens, 
1974)  for  the  lower  range  limit  of  this  species  was  based  on 
occurrence  in  the  “upper  Umpqua”  Formation  of  south- 
western Oregon.  The  “upper  Umpqua,”  however,  is  actually 
equivalent  to  both  the  “Capay”  and  “Domengine”  “Stages” 
(Baldwin,  1974;  Miles,  1981).  A lower  range  limit  of  upper- 
most “Capay,”  nevertheless,  can  be  substantiated  by  the  oc- 
currence of  this  species  in  the  zone  of  interfingering  between 
the  coastal  alluvial-fan  facies  and  the  shallow-marine  (trans- 
gressive) facies  of  the  Llajas  Formation. 

Family  Cerithiidae  Fleming,  1828 
Subfamily  Cerithiinae  Fleming,  1828 
Genus  Benoistia  Cossmann,  1899 

Type  Species.  By  original  designation,  Cerithium  muri- 
coides. 

Benoistia  umpquaensis  Turner,  1938 

Figure  6m 

Benoistia  umpquaensis  Turner,  1938:82,  pi.  21,  figs.  8,  10. 

Weaver,  1943:387,  pi.  76,  figs.  1 1-12. 

Tectanus  ligniticus  Vokes,  1939:164-165,  pi.  21,  figs.  1, 
3,  4. 

Benoistea  umpquaensis  Turner.  Givens  and  Kennedy,  1976: 
964-965,  pi.  1,  figs.  14-21. 

Primary  Type  Material.  UCMP  holotype  33192,  “upper 
Umpqua”  Formation,  UCMP  locality  A-661. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  San  Diego,  California  through 
southwestern  Oregon. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  374. 

Remarks.  Only  a single  specimen  was  found,  and  it  is  from 


the  “Stewart  bed.”  See  Givens  and  Kennedy  (1976)  for  a 
detailed  morphologic  description  of  this  species. 

The  holotype  of  Tectarius  ligniticus  is  UCMP  1 5838,  Lla- 
jas Formation,  UCMP  locality  7005. 

Family  Cerithiopsidae  H.  and  A.  Adams,  1854 
Genus  Cerithiopsis  Forbes  and  Hanley,  1853 

Type  Species.  By  monotypy,  Murex  tubercularis  Montagu, 
1803. 

Cerithiopsis  llajasensis  n.  sp. 

Figures  6n-o 

Diagnosis.  A species  of  Cerithiopsis  characterized  by  te- 
leoconch  sculpture  of  four  to  five  primary  spiral  ribs  crossed 
by  equal-strength  collabral  costae,  with  nodes  at  the  inter- 
sections. The  varices  are  irregularly  spaced. 

Cerithiopsis  llajasensis  n.  sp.  is  similar  to  C.  orovillensis 
Dickerson  (1916:489-490,  pi.  39,  fig.  7)  from  Eocene  strata 
in  northern  California.  C.  llajasensis  differs  from  the  holo- 
type of  C.  orovillensis  in  the  following  features:  larger,  a more 
elongate-slender  shape,  equal  strength  spiral  and  collabral 
sculpture,  four  to  five  rather  than  only  four  primary  spiral 
ribs  on  the  teleoconch  whorls,  equal-strength  primary  spiral 
ribs,  less  nodose  sculpture,  more  varices,  and  an  absence  of 
two  strong  spiral  keels  above  the  base  of  the  body  whorl. 

Cerithiopsis  llajasensis  n.  sp.  is  similar  also  to  C.  excelsus 
Dali  ( 1 909:75,  pi.  3,  fig.  9)  from  Eocene  strata  in  southwestern 
Oregon.  Givens  (1974)  regarded  C.  excelsus  as  C.  excelsum 
(Dali).  C.  llajasensis  differs  from  the  holotype  of  C.  excelsum 
in  the  following  features:  suture  more  adpressed,  absence  of 
a shoulder  on  the  whorls,  equally  spaced  collabral  sculpture, 
fewer  primary  spiral  ribs,  equal-strength  primary  spiral  ribs, 
many  more  varices,  and  more  swollen  varices.  The  cancellate 
sculpture  of  C.  llajasensis  is  equal  in  strength  over  the  shell 
whereas  in  C.  excelsum  this  is  not  the  case.  C.  excelsum  has 
two  prominent  spiral  ribs  and  unequally  spaced  collabral 
sculpture,  thereby  producing  an  uneven  cancellate  sculpture. 

Description.  Shell  small,  turriform  and  elongate,  with  many 
convex  whorls.  Suture  moderately  impressed,  undulating. 
Protoconch  mammillated,  about  3 Vi  whorls,  smooth.  Teleo- 
conch sculpture  cancellate,  nodes  at  intersections  of  primary 
spiral  ribs  and  collabral  costae.  Spiral  and  collabral  sculpture 
equal  strength.  Four  to  five  primary  spiral  ribs  per  whorl; 
seven  on  body  whorl  proper,  five  more  on  base  of  body  whorl. 
Ribs  equally  spaced,  usually  separated  by  a secondary  riblet. 
No  secondary  riblets  on  base  of  body  whorl.  Fourteen  to  16 
collabral  costae  per  whorl.  Costae  equally  spaced,  extending 
from  suture  to  suture.  No  collabral  ornament  on  base  of  body 
whorl.  Swollen  varices  irregularly  spaced,  usually  two  per 
whorl. 

Columella  smooth,  covered  by  a thin  callus.  Aperture  ovate. 
Anterior  canal  short,  shallowly  notched,  bent  backwards.  Di- 
mensions of  largest  specimen,  height  (incomplete)  22.5  mm, 
body  whorl  width  6 mm. 

Primary  Type  Material.  LACMIP  holotype  6515,  LAC- 
MIP  paratype  6516,  Llajas  Formation,  CSUN  locality  371. 


20  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  Simi  Valley,  California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  373,  489,  491,  498,  517. 

Remarks.  Specimens  of  this  new  species  are  abundant  at 
locality  371,  but  preservation  is  only  fair  in  most  of  the 
specimens.  The  type  specimens  have  the  best  overall  pres- 
ervation. Fifteen  specimens  were  found  at  locality  373,  one 
specimen  each  from  localities  489  and  491,  and  three  spec- 
imens each  from  localities  498  and  5 1 7.  Preservation  is  not 
as  good  at  these  other  localities  as  it  is  at  locality  37 1 . 

The  bent-backwards  anterior  canal  of  this  species  is  well 
developed  enough  to  suggest  assignment  to  Cerithiopsis.  Bit- 
tium  is  similar  to  Cerithiopsis  in  having  a short  and  shallow 
anterior  canal,  but  according  to  Houbrick  (1977)  the  canal 
is  not  bent  backwards  in  Bittium. 

Etymology.  The  species  is  named  for  the  Llajas  Formation. 


Superfamily  Epitoniacea 
Family  Epitoniidae  Lamarck,  1822 
Genus  Cirsotrema  Morch,  1852 


Type  Species.  By  monotypy,  Seal  ana  varicosum  Lamarck, 
1822. 


Cirsotrema  sp. 

Figure  6p 


Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  374. 

Remarks.  Only  a single  specimen  was  found,  and  it  is  from 
the  “Stewart  bed.”  This  genus  is  characterized  by  thick  axial 
varices  more  or  less  reflexed  posteriorly  and  usually  not  con- 
tinuous from  whorl  to  whorl;  coarse  primary  spiral  ribs;  and 
basal  disk  bounded  by  a prominent  enlargement  of  the  var- 
ices. A species  identification  could  not  be  made  as  parts  of 
the  aperture  and  basal  keel  area  are  missing. 


Superfamily  Calyptraeacea 
Family  Calyptraeidae  Blainville,  1824 
Genus  Calyptraea  Lamarck,  1799 

Type  Species.  By  monotypy.  Patella  chinensis  Linne,  1758. 

Calyptraea  diegoana  (Conrad,  1855) 

Figure  6q 

Trochita  diegoana  Conrad,  1855:7,  17;  1857:327,  pi.  5,  fig. 
42. 

Galerus  excentricus  Gabb,  1864:136,  pi.  20,  fig.  95;  pi.  29, 
fig.  232a.  Dickerson,  1913:264. 

Calyptraea  calabasaensis  Nelson,  1925:419,  pi.  54,  figs. 
8a-b. 

Calyptraea  (Galerus)  calabasaensis  Nelson.  Clark  and  Wood- 
ford, 1927:120,  pi.  21,  figs.  10-13. 

Calyptraea  diegoana  (Conrad).  Stewart,  1927:340-341,  pi. 
27,  fig.  15.  Turner,  1938:89-90,  pi.  20,  figs.  1-2.  Weaver, 
1943:351-352,  pi.  71,  figs.  16,  20;  pi.  103,  fig.  3;  1953:29. 


Stewart,  1946:pl.  11,  fig.  5.  Kleinpell  and  Weaver,  1963: 
186,  pi.  24,  fig.  7.  Hickman,  1969:79,  82,  pi.  11,  figs.  7- 
8;  1980:33-34,  pi.  2,  figs.  18-21.  Smith,  1975:469,  table 
1.  Givens  and  Kennedy,  1979:table  2. 

Primary  Type  Material.  USNM  holotype  1856,  Tejon? 
horizon,  San  Diego,  California. 

Molluscan  Stage  Range.  “Martinez”  through  lower  Oli- 
gocene. 

Geographic  Distribution.  San  Diego,  California  through 
Washington. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  455,  491,  548.  Shallow-marine  (re- 
gressive) facies:  CSUN  localities  462,  475,  51 2d. 

Remarks.  Specimens  were  most  abundant  at  locality  37  1 . 
At  all  localities,  the  specimens  are  unattached. 

Family  Xenophoridae  Philippi,  1853 

Genus  Xenophora  Fischer  von  Waldheim,  1807 

Type  Species.  By  subsequent  designation  (Gray,  1847), 
Trochus  conchyliophorus  Bom,  1780. 

Xenophora  stocki  Dickerson,  1916 

Figure  6r 

Xenophora  stocki  Dickerson,  1916:502-503,  pi.  37,  figs.  4a- 
b.  Givens,  1974:71,  pi.  7,  fig.  8.  Givens  and  Kennedy, 
1979:tables  1,  3. 

Primary  Type  Material.  UCMP  holotype  11838,  Rose 
Canyon  Shale,  UCMP  locality  2226. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  San  Diego  through  Pine  Moun- 
tain area,  southern  California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  374,  444,  458,  47  Id,  472,  477,  479. 

Remarks.  Xenophora  stocki  is  most  abundant  in  the  “Stew- 
art bed,”  especially  at  locality  374.  Specimens  at  all  localities 
occur  as  internal  molds  with  only  remnants  of  shell  material. 

Superfamily  Strombacea 
Family  Strombidae  Rafinesque,  1815 
Genus  Ectinochilus  Cossmann,  1889 

Type  Species.  By  original  designation,  Strombus  canalis. 

Subgenus  Macilentos  Clark  and  Palmer,  1923 

Type  Species.  By  original  designation,  Rimella  macilenta 
White,  1889. 

Ectinochilus  ( Macilentos ) macilentus 
(White,  1889) 

Figure  6s 

Rimella  macilenta  White,  1889:19,  pi.  3,  figs.  10-12. 
Ectinochilus  (Macilentos)  macilentus  (White).  Clark  and 
Palmer,  1923:280,  pi.  51,  figs.  9-10.  Givens,  1974:72,  pi. 
7,  figs.  13,  16.  Squires,  1977:table  1;  1983b:fig.  9b. 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  21 


Figure  6.  Llajas  Formation  gastropods.  Unless  otherwise  indicated,  views  are  apertural.  a.  Nerita  cf.  N.  (Amphinerita)  eorex  Vokes,  1939, 
LACMIP  hypotype  6508,  CSUN  loc.  700,  x 6.  b and  c.  Velates  perversus  (Gmelin,  1791),  UCMP  hypotype  37435,  UCMP  loc.  7193  = CSUN 
loc.  702,  xl.3.  b,  abapertural  view.  d.  Turritella  meganosensis  protumescens  Merriam  and  Turner,  1937,  LACMIP  6509,  CSUN  loc.  454, 
x 1.  e.  Turritella  andersoni  Dickerson,  1916,  LACMIP  hypotype  6510,  CSUN  loc.  469d,  x2.  f and  g.  Turritella  andersoni  lawsoni  Dickerson, 
1916.  All  parts  from  CSUN  loc.  374  and  x 1.25.  f,  LACMIP  hypotype  6511.  g,  LACMIP  hypotype  6512.  h.  Turritella  buwaldana  Dickerson, 


22  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Rimella  (Macilentos)  macilenta  White.  Vokes,  1939:15 5— 
156,  pi.  20,  figs.  1,  2,  4,  5. 

Ectinochilus  maci/entus  (White).  Stewart,  1946:93,  pi.  11, 
figs.  12-15.  Givens  and  Kennedy,  1979:83,  table  1. 

Primary  Type  Material.  USNM  holotype  201  14,  Domen- 
gine  Formation,  about  3 km  north  of  New  Idria,  section  16, 
T 17  S,  R 12  E,  Priest  Valley  quadrangle,  Fresno  County, 
California;  CAS  paratype  769,  Llajas  Formation,  CAS  lo- 
cality 393. 

Molluscan  Stage  Range.  “Capay”  through  “Domengine.” 
Geographic  Distribution.  San  Diego  through  central  Cal- 
ifornia. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  372,  373,  374,  440,  444,  445,  447,  449, 
455,  458,  460,  471d,  472,  481, 484,  491, 493,  494,  498,  499, 
508,  518,  538,  539.  Outer  shelf  and  slope  facies:  CSUN  lo- 
cality 482. 

Remarks.  This  species  is  common  in  the  shallow-marine 
(transgressive)  facies.  The  two  localities  at  which  it  is  most 
abundant  are  371  and  374.  At  371  specimens  are  very  well 
preserved  and  some  have  the  protoconch  or  the  fragile  outer 
lip  intact. 

The  diagnosis  of  the  subgenus  Macilentos  by  Clark  and 
Palmer  (1923)  was  based  on  the  paratype  and  other  speci- 
mens from  the  Llajas  Formation,  rather  than  on  White’s 
holotype  from  central  California. 

Family  Seraphsidae  Jung,  1974 
Genus  Paraseraphs  Jung,  1974 

Type  Species.  By  original  designation,  Paraseraphs  tetanus 
Jung,  1974  [=Terebellum  fusiforme  of  authors,  not  of  La- 
marck]. 

Paraseraphs  erraticus  (Cooper,  1894) 

Figure  7a 

Tornatina  erratica  Cooper,  1894:47,  pi.  2,  fig.  35.  Waring, 
1 9 1 7:pl.  15,  fig.  11. 

Terebellum  californicum  Vokes,  1939:157,  pi.  20,  figs.  7-8, 
1 1. 

Terebellum  ( Terebellum ) erraticum  (Cooper).  KJeinpell  and 
Weaver,  1963:189,  pi.  25,  figs.  8-9. 

Paraseraphs  erraticus  (Cooper).  Jung,  1974:41,  pi.  12,  figs. 
8-14;  pi.  13,  figs.  1-3.  Givens  and  Kennedy,  1979:87, 
tables  1,  3. 


Primary  Type  Material.  CAS  holotype  608,  Eocene  strata. 
Rose  Canyon,  San  Diego,  California. 

Molluscan  Stage  Range.  Uppermost  “Capay”  through 
“Transition.” 

Geographic  Distribution.  San  Diego  through  central  Cal- 
ifornia. 

Local  Occurrence.  Interfingering  coastal  alluvial-fan  facies 
and  shallow-marine  (transgressive)  facies:  CSUN  localities 
469d,  702.  Shallow-marine  (transgressive)  facies:  CSUN  lo- 
calities 371,  540. 

Remarks.  Only  a few  specimens  were  found,  and  most  are 
from  locality  371.  The  largest  specimen  (Fig.  7a)  is  35  mm 
in  height  and  is  nearly  twice  the  size  of  the  holotype. 

The  presence  of  P.  erraticus  in  the  zone  of  interfingering 
between  the  coastal  alluvial-fan  facies  and  the  shallow-ma- 
rine (transgressive)  facies  of  the  Llajas  Formation  extends 
the  molluscan  stage  range  of  this  taxon  into  the  uppermost 
“Capay.”  Previously,  the  lower  range  limit  had  been  reported 
as  the  “Domengine  Stage”  (Vokes,  1939;  Jung,  1974). 

Superfamily  Cypraeacea 
Family  Cypraeidae  Rafinesque,  1815 
Subfamily  Cypraeinae  Rafinesque,  1815 
Genus  Cypraea  Linne,  1758 

Type  Species.  By  subsequent  designation  (Montfort,  1810), 
Cypraea  tigris  Linne,  1758. 

Cypraea  castacensis  Stewart,  1927 

Figure  7b 

Cypraea  castacensis  Stewart,  1927:370,  pi.  28,  fig.  10.  In- 
gram, 1942:13,  pi.  1,  figs.  5-6. 

Eocypraea  castacensis  (Stewart).  Vokes,  1939:154,  pi.  20, 

figs.  9,  14. 

Primary  Type  Material.  UCMP  holotype  1 1690,  Tejon? 
Formation,  UCMP  locality  452. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  Simi  Valley  through  central  Cal- 
ifornia. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  374,  444,  491. 

Remarks.  Specimens  are  relatively  rare  in  the  Llajas  For- 


1916,  LACMIP  hypotype  6513,  CSUN  loc.  373,  x3.  i.  Turritella  uvasana  infera  Merriam,  1941,  abapertural  view,  UCLA  hypotype  59359, 
UCLA  loc.  6616,  xl.25.  j.  Turritella  uvasana  applinae  Hanna,  1927,  LACMIP  hypotype  6514,  CSUN  loc.  374,  xl.  k.  Architectonica 
(Architectonica)  llajasensis  Sutherland,  1966,  dorsal  view,  LACMIP  6515,  CSUN  loc.  371,  x3.  1.  Architectonica  ( Stellaxis ) cognata  Gabb, 
1864,  dorsal  view,  LACMIP  hypotype  6516,  CSUN  loc.  374,  x 1,  m.  Benoistia  umpquaensis  Turner,  1938,  abapertural  view,  UCLA  hypotype 
59274,  CSUN  loc.  374,  xl.  n and  o.  Cerithiopsis  llajasensis  n.  sp.  All  parts  from  CSUN  loc.  371.  n,  LACMIP  holotype  6517,  x3.5.  o, 
abapertural  view,  LACMIP  paratype  6518,  x3.  p.  Cirsotrema  sp.,  UCLA  hypotype  59275,  CSUN  loc.  374,  x 1.5.  q.  Calyptraea  diegoana 
(Conrad,  1855),  side  view,  LACMIP  hypotype  6519,  CSUN  loc.  371,  x 1.5.  r.  Xenophora  stocki  Dickerson,  1916,  internal  mold,  side  view, 
LACMIP  hypotype  6520,  CSUN  loc.  374,  xl.  s.  Ectinochilus  ( Macilentos ) macilentus  (White,  1889),  LACMIP  hypotype  6521,  CSUN  loc. 
371,  x 1.5. 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  23 


mation.  One  of  Vokes’  (1939:154,  pi.  20,  fig.  9)  hypotypes 
is  from  Devil  Canyon,  just  north  of  locality  373. 

Superfamily  Naticacea 

Family  Naticidae  Forbes,  1838 

Subfamily  Ampullospirinae  Cox,  1930 

Genus  Eocernina  Gardner  and  Bowles,  1934 

Type  Species.  By  original  designation,  Natica  hannibali 
Dickerson,  1914. 

Eocernina  hannibali  (Dickerson,  1914) 

Figure  7c 

Natica  hannibali  Dickerson,  1 9 1 4: 1 1 9,  pi.  12,  figs.  5a-b;  1916: 
508,  pi.  38,  figs.  9a-b. 

Natica  (Cryptonatica)  hannibali  Dickerson.  Waring,  1917: 
pi.  15,  figs.  21-23. 

Ampullina  hannibali  (Dickerson).  Hanna,  1927:306,  pi.  48, 
figs.  1-3,  10. 

Ampullina  (Globularia)  hannibali  (Dickerson).  Clark,  1929: 
pi.  11,  fig.  12. 

Cernina  ( Eocernina ) hannibali  (Dickerson).  Turner,  1938: 
87-88,  pi.  19,  fig.  3.  Vokes,  1939:172,  pi.  22,  figs.  1,  3. 
Weaver,  1943:348-349,  pi.  71,  figs.  8-9,  21,  23. 
Eocernina  hannibali  ( Dickerson).  Hanna  and  Hertlein,  1943: 
fig.  62-17.  Marincovich,  1977:229-231,  pi.  18,  fig.  14;  pi. 
19,  figs.  1-4.  Givens  and  Kennedy,  1979:87,  tables  1.  3. 
Squires,  1983b:fig.  9c. 

Globularia  ( Eocernina ) hannibali  (Dickerson).  Stewart,  1946: 
pi.  11,  fig.  18.  Givens,  1974:75,  pi.  9,  figs.  1,  3.  Squires, 

1 977:table  1 . 

Primary  Type  Material.  CAS  holotype  243,  Umpqua  For- 
mation, CAS  locality  25. 

Molluscan  Stage  Range.  “Capay”  through  “Transition.” 
Geographic  Distribution.  San  Diego,  California  through 
southwestern  Oregon. 

Local  Occurrence.  Interfingering  coastal  alluvial-fan  facies 
and  shallow-marine  (transgressive)  facies:  CSUN  localities 
469a,  542,  545,  702,  703.  Shallow-marine  (transgressive)  fa- 
cies: CSUN  localities  37 1 , 372,  373,  374,  439,  440,  444,  445, 
447,  449,  450,  451,  455,  458,  460,  465,  469e,  469g,  470b, 
471b,  47 le,  477,  479,  480,  484,  486,  488,  493,  494,  497, 
498,  499,  500,  508,  509,  5 10,  5 14,  5 1 7,  523,  538,  704.  Outer 
shelf  and  slope  facies:  CSUN  locality  482.  Shallow-marine 
(regressive)  facies:  CSUN  localities  512c,  51 2d. 

Remarks.  E.  hannibali  is  one  of  the  most  common  species 
of  the  Llajas  Formation.  It  occurs  in  all  the  marine  facies  of 
the  formation  and  is  particularly  characteristic  of  the  shallow- 
marine  (transgressive)  facies.  According  to  Marincovich 
( 1 977:23 1 ),  this  species  may  occur  in  prolific  numbers.  This 
is  especially  true  for  the  “Stewart  bed”  in  the  vicinity  of 
localities  374  and  444.  There,  E.  hannibali  is  the  most  com- 


mon megafossil  with  nearly  complete  growth  series  present 
also. 

Genus  Tejonia  Hanna  and  Hertlein,  1943 

Type  Species.  By  monotypy,  Natica  alveata  Conrad,  1855, 
not  Troschel,  1852. 

Tejonia  moragai  (Stewart,  1927) 

Figure  7d 

Natica  alveata  Conrad,  1855:10;  1857:321,  pi.  2,  figs.  8,8a. 
Not  Natica  alveata  Troschel,  1852:159,  pi.  5,  fig.  3. 
Amauropsis  alveata  (Conrad).  Gabb,  1864:110,  pi.  19,  fig. 
59;  pi.  2 1 , fig.  111.  Dickerson,  1 9 1 6:pl.  38,  fig.  7.  Anderson 
and  Hanna,  1925: 1 19-120,  pi.  6,  fig.  2;  pi.  7,  fig.  1;  pi.  15, 
fig.  17. 

Amauropsis  alveatus  (Conrad).  Arnold,  1 907 :pl.  39,  fig.  8. 
Amaurellina  moragai  Stewart,  1927:334-336,  pi.  28,  fig.  3 
[new  name  for  Natica  alveata  Conrad,  1855,  preoccupied]. 
Clark,  1929:pl.  14,  figs.  3,  9.  Kleinpell  and  Weaver,  1963: 
188,  pi.  25,  figs.  1-2. 

Amaurellina  moragai  lajollaensis  Stewart,  1927:336,  pi.  28, 
fig.  2. 

Amaurellina  hendoni  Turner,  1938:86-87,  pi.  20,  figs.  7,  8, 
11.  Vokes,  1939:173,  pi.  22,  figs.  7,  10.  Weaver,  1943:345, 
pi.  70,  figs.  12-13,  16-17. 

Tejonia  lajollaensis  (Stewart).  Hanna  and  Hertlein,  1 943: 
172,  fig.  62-30.  Givens,  1974:74,  pi.  8,  fig.  5. 

Tejonia  moragai  (Stewart).  Givens,  1974:74,  pi.  8,  fig.  8. 
Marincovich.  1977:232-236,  pi.  19,  figs.  6-12.  Squires, 
1977:table  1.  Givens  and  Kennedy,  1979:tables  1,  3-4. 

Primary  Type  Material.  Holotype  unknown  for  Natica 
alveata  Conrad  (USNM  collection?),  Tejon  Formation, 
Grapevine  Canyon,  Tejon  quadrangle,  Kern  County,  Cali- 
fornia. UCMP  “holotype”  31387  of  Amaurellina  moragai 
Stewart,  Tejon  Formation,  UCMP  locality  7200.  See  Marin- 
covich, 1977:235  for  a discussion  of  Stewart’s  (1927)  “ho- 
lotype.” 

Molluscan  Stage  Range.  “Domengine”  through  “Tejon.” 
Geographic  Distribution.  San  Diego,  California  through 
southwestern  Oregon. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  373,  467.  Shallow-marine  (regressive) 
facies:  CSUN  locality  489. 

Remarks.  At  locality  373,  T.  moragai  is  fairly  common 
and  occurs  as  a growth  series.  Some  specimens  have  their 
protoconchs  present.  Elsewhere,  only  single  specimens  were 
found.  This  species  is  characterized  by  its  strongly  tabulate 
adult  whorls. 

Genus  Pachycrommium  Woodring,  1928 

Type  Species.  By  original  designation,  Amaura  guppyi 
Gabb,  1873. 

Pachycrommium  clarki  (Stewart,  1927) 

Figure  7e 

Amauropsis  alveata  (Conrad).  Arnold,  1910:pl.  4,  fig.  21. 
Waring,  191 7:pl.  15,  fig.  25.  Dickerson,  191 5:pl.  5,  fig.  9 


24  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


[not  Natica  alveata  Conrad,  1855  = Tejoma  moragai 
(Stewart,  1927),  treated  herein]. 

Amaurellina  ( Euspirocrommium ) clarki  Stewart,  1927:336- 
339,  pi.  26,  figs.  8-9  [new  name,  in  part,  for  Amauropsis 
alveata  (Conrad,  1855),  preoccupied].  Clark,  1929:pl.  1 1, 
fig.  10.  Turner,  1938:86,  pi.  20,  fig.  3.  Weaver,  1943:345, 
pi.  70,  figs.  10,  18.  Kleinpell  and  Weaver,  1963:188,  pi. 
27,  fig.  15. 

Amaurellina  clarki  Stewart.  Gardner  and  Bowles,  1934:246, 
figs.  6,  8. 

Amaurellina ? multiangulata  Vokes,  1939: 1 74,  pi.  22,  figs.  2, 
8,  13. 

Pachycrommiuml  clarki  (Stewart).  Vokes,  1939:175,  pi.  22, 
figs.  11,  30.  Givens,  1974:73,  pi.  8,  figs.  6,  10.  Squires, 
1977:table  1. 

Amaurellina  ( Euspirocrommium ?)  clarki  Stewart.  Stewart, 
1946:pl.  1 1,  fig.  3. 

Pachycrommium  clarki  (Stewart).  Marincovich,  1977:238- 
241,  pi.  20,  figs.  4-10.  Squires,  1983b:fig.  9d. 

Primary  Type  Material.  UCMP  holotype  31385,  UCMP 
paratype  31386  of  Amaurellina  ( Euspirocrommium ) clarki 
Stewart,  Llajas  Formation,  UCMP  locality  7004  = CSUN 
locality  374. 

Molluscan  Stage  Range.  “Capay”  through  “Tejon. ” 
Geographic  Distribution.  San  Diego,  California  through 
northern  Washington. 

Local  Occurrence.  Interfingering  coastal  alluvial-fan  facies 
and  shallow-marine  (transgressive)  facies:  CSUN  localities 
452,  469a,  702,  703.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  37 1, 372,  373,  374,  444,  447,  455,  457,  466, 
469e,  47 le,  479,  484,  486,  488,  491,  492,  493,  494,  498, 
500,  505,  506,  507,  508,  5 1 3,  5 14,  5 1 8,  528,  537,  538.  Outer 
shelf  and  slope  facies:  CSUN  locality  482.  Outer  shelf  and 
slope  channel  facies:  CSUN  locality  541. 

Remarks.  This  species  is  especially  abundant  at  localities 
371  and  374,  and  at  both  localities  it  occurs  as  a growth 
series. 

Subfamily  Polinicinae 
Finlay  and  Marwick,  1937 

Genus  Polinices  Montfort,  1810 

Type  Species.  By  original  designation,  Polinices  albus 
Montfort,  1810. 

Subgenus  Euspira  Agassiz  in 
J.  Sowerby,  1838b 

Type  Species.  By  subsequent  designation  (Harris,  1897), 
Ampullaria  sigaretina  Lamarck,  1804. 

Polinices  ( Euspira ) nuciformis  (Gabb,  1864) 

Figure  7f 

Lunatia  nuciformis  Gabb,  1864:107,  pi.  28,  fig.  218.  Dick- 
erson, 19 1 6: pi.  39,  fig.  4. 

Lunatia  cowlitzensis  Dickerson,  1915:57,  pi.  4,  figs.  12a-b. 
Natica  nuciformis  (Gabb).  Anderson  and  Hanna,  1925:1  16, 
pi.  10,  fig.  8. 


Polinices  (Euspira)  nuciformis  (Gabb).  Clark  and  Woodford, 
1927:  121,  pi.  21,  figs.  16-17.  Turner,  1938:88,  pi.  20,  figs. 
4-5.  Clark,  1 938:703-704,  pi.  4,  figs.  26,31.  Vokes,  1 939: 

168,  pi.  21,  figs.  12-14.  Weaver,  1943:342-343,  pi.  70, 
figs.  1-2;  pi.  103,  fig.  2. 

Euspira  nuciformis  ( Gabb).  Stewart,  1927:323-324,  pi.  30, 
fig.  1 6;  1 946:pl.  1 1,  fig.  16.  Weaver,  1 953:29.  Givens,  1974: 
77,  pi.  7,  fig.  14.  Squires,  !977:table  1. 

Polinices  (Euspira)  nuciformis  var.  cowlitzensis  (Dickerson). 

Weaver,  1943:343,  pi.  69,  figs.  10-11,  13-19. 

Polinices  (Euspira)  nuciformis  (Gabb).  Marincovich,  1977: 
281-285,  pi.  26,  figs.  6-9. 

Primary  Type  Material.  ANSP  lectotype  4213,  ANSP 
paralectotypes  42 1 3 (two  specimens),  Tejon  Formation,  Live 
Oak  Canyon,  Kern  County,  California. 

Molluscan  Stage  Range.  Upper  Paleocene  through  “Te- 
jon.” 

Geographic  Distribution.  San  Diego,  California  through 
southwestern  Oregon. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  374. 

Remarks.  Only  three  specimens  were  found  in  the  Llajas 
Formation. 

Genus  Neverita  Risso,  1826 

Type  Species.  By  monotypy,  Neverita  josephina  Risso, 
1826. 

Subgenus  Neverita  s.s. 

Neverita  ( Neverita ) globosa  Gabb,  1869 

Figure  7g 

Neverita  globosa  Gabb,  1869:161,  pi.  27,  fig.  39.  Dickerson, 
1916:510,  pi.  39,  figs.  5a-b.  Stewart,  1927:326-327,  pi.  28, 
fig.  6.  Clark  and  Woodford,  1927:121-122,  pi.  22,  figs.  5- 
10.  Turner,  1938:89,  pi.  19,  figs.  6-7,  13-15.  Vokes,  1939: 

169,  pi.  21,  figs.  9,  15-19.  Givens  and  Kennedy,  1979: 
tables  1-3. 

Neverita  weaver i Dickerson,  1915:57,  pi.  4,  figs.  lOa-b. 
Neverita  nomlandi  Dickerson,  1917:173-174,  pi.  30,  figs. 
2a-b. 

Polinices  weaven  (Dickerson).  Turner,  1938:86,  pi.  20,  figs. 
14,  16. 

Neverita  globosa  reefensis  Vokes,  1 939: 1 69,  pi.  21,  figs.  24- 
25. 

Polinices  (Neverita)  weaveri  (Dickerson).  Weaver,  1943:340, 
pi.  68,  figs.  16-17;  pi.  69,  fig.  3. 

Polinices  (Neverita)  globosa  (Gabb).  Weaver,  1943:339,  pi. 

68,  figs.  21,  24;  pi.  69,  figs.  5-6;  pi.  100,  fig.  29. 

Polinices  (Neverita)  nomlandi  (Dickerson).  Weaver,  1943: 
340,  pi.  69,  figs.  8,  9,  12. 

Neverita  (Neverita)  globosa  Gabb.  Givens,  1974:76.  Marm- 
covich,  1977:312-316,  pi.  28,  figs.  10-15;  pi.  29,  figs. 
1-3. 

Neverita  (Glossaulaxl)  globosa  Gabb.  Givens  and  Kennedy, 
1976:965-966,  pi.  2,  figs.  5-14,  16,  18-19. 

Neverita  globosa  Gabb.  Givens  and  Kennedy,  1979:tables 
1-3. 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  25 


Primary  Type  Material.  MCZ  holotype  27859,  Domen- 
gine?  Formation,  1 6 km  west  of  Griswold’s,  on  the  road  from 
San  Juan  to  New  Idria,  and  southeast  of  the  “Sheep  Well,” 
T 15  S,  R 9 E,  Priest  Valley  quadrangle,  San  Benito  County, 
California. 

Molluscan  Stage  Range.  “Meganos”  through  upper  Eocene. 
Geographic  Distribution.  San  Diego,  California  through 
western  Washington. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  704. 

Remarks.  Only  three  specimens  were  found  in  the  Llajas 
Formation. 

Subfamily  Sininae  Wenz,  1941 
Genus  Sinum  Roding,  1798 

Type  Species.  By  subsequent  designation  (Dali,  1915),  He- 
lix  haliotoidea  Linne,  1758. 

Simon  obliquum  (Gabb,  1 864) 

Figure  7h 

Naticina  obliqua  Gabb,  1 864: 1 09,  pi.  21,  fig.  11 2.  Dickerson, 
1 9 1 5 :pl.  5,  figs.  5a-b. 

Simon  occidentis  Weaver  and  Palmer,  1922:32-33,  pi.  11, 
hgs.  8,  26.  Weaver,  1943:351,  pi.  71,  fig.  15. 

Sinum  coryliforme  Anderson  and  Hanna,  1925:120,  pi.  9, 
fig.  10;  pi.  10,  fig.  15;  pi.  15,  fig.  8. 

Sinum  obliquum  (Gabb).  Stewart,  1927:327,  pi.  30,  fig.  7a. 
Clark,  1938:704,  pi.  3,  figs.  32,  37.  Weaver,  1 943:350- 
351,  pi.  71,  fig.  13;  pi.  103,  fig.  6.  Hickman,  1969:85-88, 
pi.  11,  figs.  9-10.  Marincovich,  1977:347-350,  pi.  33,  figs. 
1-12.  Squires,  1977:table  1.  Givens  and  Kennedy,  1979: 
table  4. 

Primary  Type  Material.  ANSP  lectotype  4215,  ANSP 
paralectotypes  4215  (seven  specimens),  Tejon  Formation, 
Fort  Tejon  area,  Kern  County,  California. 

Molluscan  Stage  Range.  “Capay”  through  lower  Oligo- 
cene. 

Geographic  Distribution.  San  Diego,  California  through 
southwestern  Washington. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  539. 

Remarks.  S',  obliquum  is  most  common  at  locality  371 
where  it  is  well  preserved  and  occurs  as  a growth  series.  At 
locality  539,  only  a single  specimen  was  found. 

Subfamily  Naticinae  Forbes,  1838 

Genus  Natica  Scopoli,  1777 

Type  Species.  By  subsequent  designation  (Anton,  1839), 
Nerita  vitellus  Linne,  1758. 

Subgenus  Naticarius  Dumeril,  1806 

Type  Species.  By  monotypy,  Nerita  canrena  Linne,  1758. 


Natica  ( Naticarius ) aff. 

N.  ( N .)  uvasana  Gabb,  1864 

Figure  7i 

Local  Occurrence.  Interfingering  coastal  alluvial-fan  facies 
and  shallow-marine  (transgressive)  facies:  CSUN  locality  545. 
Shallow-marine  (transgressive)  facies:  CSUN  localities  371, 
373,  486. 

Remarks.  This  taxon  is  most  common  at  locality  37 1 where 
32  specimens  make  up  a growth  series.  The  largest  specimen 
is  18  mm  in  height.  Elsewhere,  only  a few  specimens  were 
found. 

The  Llajas  specimens  agree  with  the  description  of  N.  ( N .) 
uvasana  given  by  Marincovich  (1977:390)  in  possessing  ir- 
regularly spaced  collabral  wrinkles  immediately  below  the 
suture,  an  open  umbilicus,  and  a robust  umbilical  callus  and 
funicle.  The  Llajas  specimens  differ  in  that  they  lack  a parietal 
callus. 

Superfamily  Tonnacea 
Family  Cassididae  Latreille,  1825 
Genus  Galeodea  Link,  1 807 

Type  Species.  By  monotypy,  Buccinum  echinophorum 
Linne,  1758. 

Subgenus  Caliagaleodea  Clark,  1 942 

Type  Species.  By  original  designation,  Caliagaleodea  cal- 
ifornica  C lark,  1942. 

Galeodea  ( Caliagaleodea ) californica 
Clark,  1942 
Figure  7j 

Galeodea  ( Caliagaleodea ) californica  Clark,  1942:1  18-1 19, 

pi.  19,  figs.  15-19. 

Galeodea  californica  Clark.  Givens  and  Kennedy,  1979: 

table  1. 

Primary  Type  Material.  UCMP  holotype  34376,  UCMP 
paratype  34377,  Llajas  Formation,  UCMP  locality  7004  = 
CSUN  locality  374. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  San  Diego  through  Simi  Valley, 
California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  374,  444,  445,  447,  458. 

Remarks.  Nearly  all  the  specimens  of  this  species  were 
found  at  the  type  locality.  The  14  specimens  at  locality  374 
make  up  a nearly  complete  growth  series,  but  most  occur  as 
internal  molds.  A few  other  specimens  were  found  elsewhere 
in  the  “Stewart  bed”  in  the  vicinity  of  locality  374. 

Subgenus  Gomphopages  Gardner,  1939 

Type  Species.  By  original  designation,  Galeodea  turneri 
Gardner,  1939. 


26  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Galeodea  ( Gomphopages ) susanae  Schenck,  1926 

Figure  7k 

Galeodea  susanae  Schenck,  1926:85,  pi.  15,  figs.  3-7.  Turner, 
1938:92,  pi.  18,  fig.  18.  Weaver,  1943:402-403,  pi.  78, 
figs.  2-3. 

Galeodea  ( Gomphopages ) susanae  Schenck.  Durham,  1942: 
184.  Givens,  1974:78. 

Primary  Type  Material.  CAS  holotype  1753,  CAS  para- 
type  1754,  Llajas  Formation,  CAS  locality  372;  CAS  para- 
type  1755,  Llajas  Formation,  CAS  locality  364;  CAS  para- 
type  1756,  near  Roseburg,  Oregon. 

Molluscan  Stage  Range.  “Domengine,”  “Transition”?. 
Geographic  Distribution.  Simi  Valley,  California  through 
southwestern  Oregon. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  371. 

Remarks.  The  16  specimens  from  locality  371  make  up  a 
growth  series,  and  they  are  extremely  well  preserved.  There 
is  excellent  preservation  of  spines  and,  in  a few  cases,  the 
long  anterior  canal.  The  largest  specimen  is  53  mm  in  height. 

Genus  Phalium  Link,  1807 

Type  Species.  By  subsequent  designation  (Dali,  1909), 
Buccinum  glaucum  Linne,  1758. 

Subgenus  Semicassis  Morch,  1852 

Type  Species.  By  subsequent  designation  (Flarris,  1897), 
Cassis  japonica. 

Phalium  ( Semicassis ) tuberculiformis 
(Hanna,  1924) 

Figure  71 

Mono  (Sconsia)  tuberculatus  Gabb,  1864:104,  pi.  19,  fig.  57. 

Arnold,  1907:pl.  39,  fig.  9. 

Not  Cassidaria  tuberculata  Risso,  1826:186. 

Morio  tuberculatus  Gabb.  Dickerson,  1913:264. 

Galeodea  tuberculata  (Gabb).  Dickerson,  191 6:pl.  42,  fig.  2. 
Galeodea  (Morio)  tuberculata  (Gabb).  Waring,  1 9 1 7:pl.  15, 
fig.  17. 

Galeodea  tuberculiformis  Hanna,  1924:167  [new  name  for 
Morio  ( Sconsia ) tuberculatus  Gabb,  1864,  preoccupied]. 
Schenck,  1926:83-84,  pi.  14,  figs.  12-16.  Stewart,  1927: 
380-38 1 , pi.  28,  fig.  11.  Vokes,  1 939: 1 49-1 50,  pi.  1 9,  figs. 
19,  21,  23-27. 

Coalingodea  tuberculiformis  (Hanna).  Durham,  1942:186,  pi. 
29,  figs.  5,  9.  Givens,  1974:78-79,  pi.  8,  fig.  7.  Squires, 
1977:table  1. 

Cassis  (Coalingodea)  tuberculata  (Gabb).  Abbott,  1968b:  59- 
60,  pi.  34. 

Phalium  (Semicassis)  tuberculiformis  (Hanna).  Givens  and 
Kennedy,  1979:82,  95,  tables  1,  3. 

Primary  Type  Material.  ANSP  lectotype  4343  of  Morio 
(Sconsia)  tuberculatus  Gabb  and  Galeodea  tuberculiformis 
Hanna,  Tejon?  Formation,  Martinez,  California. 


Molluscan  Stage  Range.  “Capay”?,  “Domengine”  through 
“Transition.” 

Geographic  Distribution.  San  Diego,  California  through 
northwestern  Washington. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  372,  373,  374,  440,  444,  445,  455,  458, 
471b,  480,  488,  494,  514,  537.  Outer  shelf  and  slope  channel 
facies:  CSUN  locality  541.  Shallow-marine  (regressive)  fa- 
cies: CSUN  localities  462,  51 2d. 

Remarks.  This  taxon  is  most  common  and  best  preserved 
at  localities  371  and  374,  where  it  occurs  in  growth  series. 
The  beaded  appearance  of  the  whorls  and  the  toothed  outer 
lip  are  well  preserved. 

Dickerson  (1913)  reported  this  species  from  “Capay”-age 
strata  in  central  California,  but  he  gave  no  specific  locality 
information  nor  did  he  figure  the  species.  For  these  reasons, 
the  “Capay”-age  occurrence  is  uncertain. 

The  presence  of  P.  (S.)  tuberculiformis  in  the  Llajas  For- 
mation is  one  of  the  earliest  occurrences  of  Phalium  on  the 
West  Coast.  It  is  also  known  from  similar  age  strata  in  San 
Diego  County  (Givens  and  Kennedy,  1979). 

Family  Cymatiidae  Iredale,  1913 
Genus  Cymatium  Roding,  1798 

Type  Species.  By  subsequent  designation  (Dali,  1 904),  Mu- 
rex  femorale  Linne,  1758. 

Subgenus  Septa  Perry,  1810 

Type  Species.  By  monotypy,  Septa  scarlatina  Perry,  1810 
(=Murex  rubecula  Linne,  1758). 

Cymatium  {Septa)  janetae  Squires,  1983a 

Figure  7 m 

Cymatium  (Septa)  janetae  Squires,  1983a:355-357,  figs. 

2a-d. 

Primary  Type  Material.  UCLA  holotype  59191,  Llajas 
Formation,  CSUN  locality  444;  UCLA  paratype  59192,  Lla- 
jas Formation,  CSUN  locality  445;  UCLA  paratype  59193, 
Llajas  Formation,  CSUN  locality  371. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  Simi  Valley,  California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  374,  444,  498. 

Remarks.  Only  juvenile  specimens  occur  at  localities  371 
and  498.  Only  adult  specimens  occur  in  the  “Stewart  bed” 
at  and  near  localities  374  and  444.  The  figured  specimen 
(Fig.  7m)  is  the  holotype.  C.  (S’.)  janetae  is,  at  present,  the 
earliest  species  worldwide  referable  to  Septa  (Squires,  1 983a). 

Genus  Ranella  Lamarck,  1816 

Type  Species.  By  subsequent  designation  (Children,  1 823), 
Ranella  gigantea  Lamarck,  1816  [=Murex  o/earium  Linne, 
1758], 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  27 


Figure  7.  Llajas  Formation  gastropods  (continued).  Unless  otherwise  indicated,  views  are  apertural.  a.  Paraseraphs  erraticus  (Cooper,  1894), 
abapertural  view,  LACM1P  hypotype  6522,  CSUN  loc.  371,  x 1.  b.  Cypraea  castacensis  Stewart,  1927,  LACM1P  hypotype  6523,  CSUN  loc. 
374,  x 1.25.  c.  Eocermna  hannibali  (Dickerson,  1914),  hypotype  LACMIP  6524,  CSUN  loc.  374,  x 1.  d.  Tejonia  moragai  (Stewart,  1927), 
LACMIP  hypotype  6525,  CSUN  loc.  467,  x l.  e.  Pachycrommium  clarki  (Stewart,  1927),  LACMIP  hypotype  6526,  CSUN  loc.  371,  x 1.  f. 
Polinices  ( Euspira ) nuciformis  (Gabb,  1864),  UCLA  hypotype  59276,  CSUN  loc.  374,  x 1 . g.  Neverita  (N event  a)  globosa  Gabb,  1 869,  LACMIP 
hypotype  6527,  CSUN  loc.  704,  x 1.5.  h.  Sinum  obliquum  (Gabb,  1864),  side  view,  LACMIP  hypotype  6528,  CSUN  loc.  371,  x2.  i.  Natica 


28  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Ranella  katherineae  Squires,  1983a 

Figure  7n 

“ Ranella ” sp.  Smith,  1970:523. 

Ranella  katherineae  Squires,  1983a:357-359,  figs.  2e-g. 

Primary  Type  Material.  UCLA  holotype  45969,  Llajas 
Formation,  CSUN  locality  374. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  Simi  Valley,  California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  374. 

Remarks.  Only  a single,  nearly  complete  adult  specimen 
was  found  in  the  Llajas  Formation.  This  specimen  is  the 
holotype  and  the  figured  specimen  (Fig.  7n).  R.  katherineae 
is  one  of  the  earliest  West  Coast  species  of  Ranella  s.s.  (Squires, 
1983a). 

Family  Bursidae  Thiele,  1925 
Genus  Olequahia  Stewart,  1927 

Type  Species.  By  original  designation,  Cassidaria  wash- 
ingtoniana  Weaver,  1912. 

Olequahia  domenginica  (Vokes,  1939) 

Figure  7o 

Ranella  domenginica  Vokes,  1939:147-148,  pi.  19,  figs.  6, 
20. 

Olequahia  hornii  domenginica  (Vokes).  Stewart,  1946: 
table  1. 

Olequahia  domenginica  (Vokes).  Givens,  1974:80,  pi.  9,  figs. 
4-5.  Givens  and  Kennedy,  1979:table  1. 

Primary  Type  Material.  UCMP  syntypes  1 5803  and  1 5804, 
Domengine  Formation,  UCMP  locality  672. 

Molluscan  Stage  Range.  “Domengine  Stage.” 

Geographic  Distribution.  San  Diego  through  central  Cal- 
ifornia. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  374,  444,  445,  47 Id,  494. 

Remarks.  Specimens  are  most  common  in  the  “Stewart 
bed”  at  locality  374  where  they  form  a growth  series. 

Vokes  (1939:147-148)  used  non-type,  supplementary 
specimens  from  the  vicinity  of  locality  374  to  augment  his 
description  of  this  taxon. 

Genus  Ranellina  Conrad,  1865 

Type  Species.  By  original  designation,  Ranellina  maclurii 
Conrad,  1865. 


Ranellina  pilsbryi  Stewart,  1927 

Figure  7p 

Not  Clavatulal  californica  Conrad,  1855:11;  1857:322,  pi. 
2,  fig.  11. 

Fusus  californicus  (Conrad).  Gabb,  1864:85-86,  pi.  28,  figs. 
205-205a. 

Nyctilochus  hornii  (Gabb).  Dickerson,  1915 :pl.  7,  fig.  8.  Dick- 
erson, 1916:pl.  42,  fig.  3. 

Clavilithes  californicus  (Conrad).  Anderson  and  Hanna,  1925: 
63-64,  pi.  13,  figs.  1-2. 

Ranellina  pilsbryi  Stewart,  1927:384-385,  pi.  30,  figs.  8-9 
[new  name  for  Fusus  californicus  (Conrad)  Gabb,  1864, 
misidentified].  Turner,  1938:91,  pi.  16,  fig.  3.  Vokes,  1939: 
148,  pi.  19,  figs.  10,  17.  Weaver,  1943:417,  pi.  82,  fig.  7. 
Givens,  1974:80,  pi.  9,  fig.  12.  Givens  and  Kennedy,  1979: 
tables  1,4. 

Primary  Type  Material.  UCMP  holotype  31382,  UCMP 
paratype  31383  of  Ranellina  pilsbryi  Stewart,  Tejon  For- 
mation, UCMP  locality  7182. 

Molluscan  Stage  Range.  “Domengine”  through  “Tejon.” 
Geographic  Distribution.  San  Diego,  California  to  south- 
western Oregon. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  373,  486,  498. 

Remarks.  This  species  is  most  common  at  locality  371 
where  seven  specimens  were  found. 

Smith  (1975:table  1)  reported  “ Ranellina ” sp.  afif.  R.  pils- 
bryi from  “Martinez”-age  strata  of  central  California. 

Family  Ficidae  Conrad,  1867 

Genus  Ficopsis  Conrad,  1866 

Type  Species.  By  subsequent  designation  (Stewart,  1927), 
Hemifusus  remondii  Gabb,  1864. 

Ficopsis  cooperiana  Stewart,  1927 

Figure  8a 

Fusus  (Hemifusus)  cooperii  Gabb,  1864:86,  pi.  28,  fig.  207. 
Not  Fusus  cooperi  Conrad,  1834:148. 

Ficopsis  cooperii  (Gabb).  Dickerson,  1915:61-62,  pi.  6,  fig. 

11;  1916:492-493,  pi.  37,  fig.  7.  Clark,  1929:pl.  9,  fig.  13. 
Ficopsis  cooperiana  Stewart,  1927:378-379  [new  name  for 
Fusus  (Hemifusus)  cooperii  Gabb,  1864,  preoccupied]. 
Clark,  1929:pl.  9,  fig.  13.  Givens,  1974:81,  pi.  9,  figs.  7, 
9.  Givens  and  Kennedy,  1979:87,  table  3. 

Primary  Type  Material.  UCMP  holotype  11691  of  Fusus 
(Hemifusus)  cooperii  Gabb  and  Ficopsis  cooperiana  Stewart, 
Eocene  strata,  UCMP  locality  2226. 


(Naticarius)  aff.  N.  (N.)  uvasana  Gabb,  1864,  LACMIP  hypotype  6529,  CSUN  loc.  371,  x 2.  j.  Galeodea  (Caliagaleodea)  californica  Clark, 
1942,  internal  mold,  abapertural  view,  LACMIP  topotype  and  hypotype  6530,  CSUN  loc.  374,  xl.  k.  Galeodea  (Gomphopages)  susanae 
Schenck,  1926,  LACMIP  hypotype  6531,  CSUN  loc.  371,  x 1.5.  1.  Phalium  (Semicassis)  tuberculiformis  (Hanna,  1924),  LACMIP  hypotype 
6532,  CSUN  loc.  371,  xl.  m.  Cymatium  (Septa)  janetae  Squires,  1983a,  UCLA  holotype  59191,  CSUN  loc.  444,  xl.  n.  Ranella  katherineae 
Squires,  1983a,  UCLA  holotype  45969,  CSUN  loc.  374,  x0.5.  o.  Olequahia  domenginica  (Vokes,  1939),  LACMIP  hypotype  6533,  CSUN 
loc.  374,  x l.  p.  Ranellina  pilsbryi  Stewart,  1927,  LACMIP  hypotype  6534,  CSUN  loc.  371,  x2.5. 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  29 


Molluscan  Stage  Range.  “Domengine”  through  “Transi- 
tion.” 

Geographic  Distribution.  San  Diego  through  central  Cal- 
ifornia. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  374. 

Remarks.  Only  a few  specimens  were  found  in  the  Llajas 
Formation. 

Ficopsis  remondii  crescentensis 
Weaver  and  Palmer,  1922 

Figure  8 b 

Ficopsis  angulatus  Weaver,  1905:1  19,  pi.  13,  fig.  5. 

Not  Pyrula  angiilata  Edwards,  1866:pl.  4. 

Ficopsis  remondii  (Gabb)  var.  crescentensis  Weaver  and 
Palmer,  1922:39-40,  pi.  11,  fig.  14  [new  name  for  Ficopsis 
angulatus  Weaver,  1 905,  preoccupied],  Stewart,  1930:40- 
41.  Turner,  1938:93,  pi.  15,  fig.  19.  Weaver,  1943:399,  pi. 
77,  fig.  10. 

Ficopsis  remondii  crescentensis  Weaver  and  Palmer.  Vokes, 
1939:152-153.  Givens,  1974:82,  pi.  9,  fig.  11.  Squires, 
1977:table  1.  Givens  and  Kennedy,  1979:87,  tables  1,  3. 
Ficopsis  crescentensis  Weaver  and  Palmer.  Stewart,  1946:pl. 
1 1,  fig.  17. 

Primary  Type  Material.  UCMP  holotype  1 1887  of  Ficop- 
sis angulatus  Weaver,  Eocene  strata,  UCMP  locality  337. 
UW  holotype  205  (CAS  7616)  of  Ficopsis  remondii  crescen- 
tensis Weaver  and  Palmer,  Crescent?  Formation,  UW  local- 
ity 358. 

Molluscan  Stage  Range.  “Capay”  through  “Transition.” 
Geographic  Distribution.  San  Diego,  California  through 
northwestern  Washington. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  373,  374,  477,  479,  486. 

Remarks.  Specimens  are  most  common  at  locality  371 
although  the  largest  specimens  occur  at  locality  374.  This 
taxon  is  characterized  by  the  tri-carination  of  the  body  whorl 
and  the  cancellate  sculpture. 

Order  Neogastropoda 

Superfamily  Muricacea 

Family  Muricidae  da  Costa,  1776 

Subfamily  Muricinae  da  Costa,  1776 

Genus  Laevityphis  Cossmann,  1903 

Type  Species.  By  original  designation,  Typhis  coronarius 
Deshayes,  1865. 

Subgenus  Laevityphis  s.s. 

Laevityphis  ( Laevityphis ) antiquus 
(Gabb,  1864) 

Figure  8c 

Typhis  antiquus  Gabb,  1864:82,  pi.  18,  fig.  31.  Stewart,  1927: 
387-388,  pi.  27,  figs.  7-8. 


Laevityphis  ( Laevityphis ) antiquus  (Gabb).  Keen,  1944:58, 

63.  Givens,  1974:82. 

Laevityphis  antiquus  (Gabb).  Givens  and  Kennedy,  1979: 

table  3. 

Primary  Type  Material.  ANSP  lectotype  4335,  Tejon? 
Formation,  Martinez,  California. 

Molluscan  Stage  Range.  “Domengine”  through  “Tejon.” 

Geographic  Distribution.  San  Diego  through  central  Cal- 
ifornia. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  371. 

Remarks.  Only  a single  specimen  was  found  in  the  Llajas 
Formation. 

Family  Buccinidae  Rafinesque,  1815 
Genus  Ancistrolepis  Dali,  1895 

Type  Species.  By  original  designation,  Chrysodomus  eu- 
cosmius  Dali,  1891. 

Ancistrolepis ? carolineae  n.  sp. 

Figures  8d-e 

Diagnosis.  Bucciniform  shell  with  two  spiral  ribs  on  spire 
and  about  six  spiral  ribs  on  body  whorl,  interspaces  char- 
acteristically concave  with  numerous  fine  lirae.  Due  to  the 
fact  that  the  canal  area  has  been  broken  off  on  each  of  the 
Llajas  specimens,  positive  generic  assignment  cannot  be  made. 

Ancistrolepis' ? carolineae  n.  sp.  is  very  similar  to  A.  clarki 
Tegland  (1933: 1 3 1-132,  pi.  12,  fig.  14)  and  A.  clarki  variety? 
Tegland  (1933:132,  pi.  12,  figs.  15-17)  from  the  Oligocene 
Blakeley  Formation,  northwestern  Washington.  A.1  caroli- 
neae differs  from  A.  clarki  and  A.  clarki  var.?  in  the  following 
features:  less  convex  whorls,  narrower  and  shorter  spire, 
equally  spaced  primary  spiral  ribs  on  the  spire,  less  inflated 
spiral  ribs,  and  smaller  shell  size.  Unfortunately,  the  anterior 
canal  area  is  missing  in  the  Llajas  specimens  and  comparison 
to  the  ornamented  canal  areas  of  A.  clarki  and  A.  clarki  var.? 
cannot  be  made.  A.  clarki  and  A.  clarki  var.?  differ  from  each 
other  only  in  the  nature  of  the  canal  area  ornamentation. 

Description.  Medium  shell,  bucciniform,  with  five  or  six 
angulate  whorls.  Suture  moderately  impressed.  Protoconch 
smooth  with  low,  rounded  whorls  and  shallow  sutures. 

Spire  whorls  with  two  very  prominent,  primary  spiral  ribs 
that  divide  the  whorls  into  three  equal  concave  areas  covered 
with  numerous  fine  lirae.  Body  whorl  with  five  to  six  primary 
spiral  ribs,  interspaces  concave  and  covered  with  numerous 
fine  lirae. 

Columella  smooth  and  twisted.  Aperture  straight.  Thick- 
ened outer  lip.  Anterior  canal  area  missing.  Dimensions  of 
largest  specimen  (Figs.  8d-e):  height  (incomplete)  32  mm, 
body  whorl  width  20  mm. 

Primary  Type  Material.  UCLA  holotype  59401,  UCLA 
paratypes  59402  (three  specimens),  Llajas  Formation,  CSUN 
locality  374. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  Simi  Valley,  California. 


30  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  374. 

Remarks.  Specimens  of  this  new  species  were  found  only 
at  CSUN  locality  374  (“Stewart  bed”).  A total  of  12  speci- 
mens was  found,  and  preservation  is  mostly  as  internal  molds. 
A few  specimens  (i.e.,  the  primary  type  material)  have  por- 
tions of  the  original  shell  material. 

Ancistrolepis  Dali  is  differentiated  from  Chrvsodomus  in 
three  respects;  namely,  a shorter  canal,  a degenerate  radula, 
and  in  the  nature  of  the  operculum.  Although  only  the  canal 
feature  can  be  used  in  fossil  material,  Tegland  (1933)  con- 
sidered that  the  outline  of  the  shell  and  detail  of  sculpturing 
of  A.  clarki  was  so  close  to  the  Recent  species,  Ancistrolepis 
eucosimus  Dali  (1895:709,  pi.  29,  fig.  7),  the  type  species  of 
the  genus,  that  reference  to  that  genus  rather  than  Chryso- 
domus  was  warranted.  The  Llajas  specimens  of  A.?  carolineae 
n.  sp.  are  very  similar  to  those  of  A.  clarki  and  A.  clarki  var.?, 
and  they  are  not  similar  to  West  Coast  Paleogene  species  of 
Chrysodomus  in  terms  of  sculpture. 

If  A.1  carolineae  does  belong  to  Ancistrolepis,  it  would 
extend  the  earliest  occurrence  of  this  genus  to  the  early  middle 
Eocene,  and  A.  carolineae  would  be  the  oldest  known  species 
referable  to  this  genus.  It  would  be  the  first  occurrence  of 
this  genus  in  California. 

Etymology.  The  species  is  named  for  Caroline  Squires. 

Superfamily  Buccinacea 
Family  Nassariidae  Iredale,  1916 
Genus  Mo/opophorus  Gabb,  1869 

Type  Species.  By  monotypy,  Bullia  (Molopophorus)  striata 

Gabb,  1869. 

Molopophorus  cretaceus  (Gabb,  1 864) 

Figure  8f 

Nassa  cretacea  Gabb,  1864:97,  pi.  18,  fig.  49. 
Molopophorus  cretaceus  ( Gabb).  Stewart,  1927:391-392,  pi. 

28,  fig.  9.  Vokes,  1939:141-142,  pi.  19,  fig.  4. 

Primary  Type  Material.  ANSP  lectotype  4197,  Domen- 
gine  Formation,  Bulls  Head  Point,  near  Martinez,  California. 

Mollusean  Stage  Range.  “Domengine.” 

Geographic  Distribution.  Simi  Valley  through  central  Cal- 
ifornia. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  467. 

Remarks.  Only  two  specimens  were  found  in  the  Llajas 
Formation,  and  the  apertures  are  covered  or  smashed. 

Family  Fasciolariidae  Gray,  1853 
Genus  Clavilithes  Swainson,  1840 

Type  Species.  By  subsequent  designation  (Grabau,  1904), 
Fusus parisiensis  Mayer-Eymar,  1877  [=Fusus  longaevus  La- 
marck, 1803,  not  Solander,  1766], 


Clavilithes  tabulatus  (Dickerson,  1913) 

Figure  8g 

Clavella  tabulata  Dickerson,  1913:283,  pi.  12,  fig.  7. 
Clavilithes  tabulatus  (Dickerson).  Clark  and  Vokes,  1936: 
874,  pi.  1,  fig.  3 (holotype  refigured).  Givens,  1974:85,  pi. 
10,  figs.  4-5. 

Clavilithes  cf.  C.  tabulatus  (Dickerson).  Crowell  and  Susuki, 
1959:588-589,  pi.  2,  figs.  6-7. 

Primary  Type  Material.  UCMP  holotype  1 1753,  Capay 
Formation,  UCMP  locality  1853. 

Mollusean  Stage  Range.  “Capay”  through  "Domengine.” 
Geographic  Distribution.  Orocopia  Mountains,  southern 
California  through  central  California. 

Local  Occurrence.  Interfingering  coastal  alluvial-fan  facies 
and  shallow-marine  (transgressive)  facies:  CSUN  locality  703. 
Shallow-marine  (transgressive)  facies:  CSUN  localities  371, 
374,  704. 

Remarks.  Only  two  specimens  were  found.  The  presence 
of  C.  tabulatus  in  the  middle  part  of  the  Llajas  extends  the 
mollusean  stage  range  of  this  taxon  into  the  “Domengine 
Stage.”  Previously  it  had  been  reported  as  confined  to  the 
“Capay  Stage”  (Clark  and  Vokes,  1936;  Crowell  and  Susuki, 
1959;  Givens,  1974). 

Clavilithes  n.  sp.  A Clark  and  Yokes,  1936 

Figure  8h 

Clavilithes  n.  sp.  Clark  and  Vokes,  1936:874,  pi.  1,  fig.  1. 
Givens  and  Kennedy,  1976:973,  pi.  4,  figs.  9,  12. 

Type  Material.  UCMP  hypotype  15468,  Llajas  Forma- 
tion, UCMP  locality  3296  (Clark  and  Vokes,  1936).  UCR 
hypotype  4865/13,  middle  Eocene  strata,  UCR  locality  4865 
(Givens  and  Kennedy,  1976).  LACMIP  hypotype  6540,  Lla- 
jas Formation,  CSUN  locality  704. 

Mollusean  Stage  Range.  “Domengine.” 

Geographic  Distribution.  San  Diego  through  Simi  Valley, 
California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  704. 

Remarks.  Two  specimens  of  this  taxon  were  found  with 
C.  tabulatus  at  CSUN  locality  704.  C.  n.  sp.  A is  character- 
ized by  the  presence  of  nodes  on  the  spire.  It  is  not  named 
at  this  time  pending  further  taxonomic  research  by  Jack 
Mount,  Rutgers  University. 

Clavilithes  n.  sp.  B Squires,  1983a 

Figure  8i 

Clavilithes  n.  sp.  Squires,  1983a:359,  figs.  2h-i. 

Type  Material.  UCLA  hypotype  59194,  Llajas  Formation, 
CSUN  locality  445;  UCLA  hypotype  59195,  Llajas  For- 
mation, CSUN  locality  444. 

Mollusean  Stage  Range.  “Domengine.” 

Geographic  Distribution.  Simi  Valley,  California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  374,  444,  445. 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  31 


Figure  8.  Llajas  Formation  gastropods  (continued).  Unless  otherwise  indicated,  views  are  apertural.  a.  Ficopsis  cooperiana  Stewart,  1927, 
side  view,  UCLA  hypotype  59277,  CSUN  loc.  374,  x 2.  b.  Ficopsis  remondii  crescentensis  Weaver  and  Palmer,  1922,  LACMIP  hypotype 
6535,  CSUN  loc.  374,  xl.  c.  Laevityphis  ( Laevityphis ) antiquus  (Gabb,  1864),  LACMIP  hypotype  6536,  CSUN  loc.  371,  x2.  d and  e. 
Ancistrolepisl  carolineae  n.  sp.  All  parts  UCLA  holotype  5940 1 , CSUN  loc.  374,  x 1 .25.  e,  abapertural  view.  f.  Molopophorus  cretaceus  (Gabb, 
1864),  abapertural  view,  LACMIP  hypotype  6537,  CSUN  loc.  467,  x7.  g.  Clavilithes  tabulatus  (Dickerson,  1913),  LACMIP  hypotype  6538, 
CSUN  loc.  371,  xQ.5.  h.  Clavilithes  n.  sp.  A Clark  and  Yokes,  1936,  LACMIP  hypotype  6539,  CSUN  loc.  704,  x 1.25.  i.  Clavilithes  n.  sp.  B 


32  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Remarks.  This  taxon  occurs  with  C.  tabulatus  at  CSUN 
locality  374.  C.  n.  sp.  B differs  from  C.  n.  sp.  A in  the  presence 
of  an  overhanging  carina  and  in  the  lack  of  nodes  on  the 
spire.  C.  n.  sp.  B is  not  named  at  this  time  pending  further 
taxonomic  research  by  Jack  Mount,  Rutgers  University. 

Subfamily  Fusininae  Swainson,  1 840 
Genus  Fusinus  Rafinesque,  1815 

Type  Species.  By  monotypy,  Murex  co/us  Linne,  1758. 

Fusinus  teglandae  Hanna,  1927 

Figure  8j 

Fusinus  teglandi  Hanna,  1927:315,  pi.  51,  fig.  9. 

Fusinus  teglandae  Hanna.  Givens,  1974:85,  pi.  10,  figs. 

6-7. 

Primary  Type  Material.  UCMP  holotype  31 124,  Ardath 
Shale,  UCMP  locality  5062. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  San  Diego  through  Pine  Moun- 
tain, southern  California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  374. 

Remarks.  Only  three  poorly  preserved  specimens  were 
found  in  the  Llajas  Formation.  The  upper  spire  is  missing 
in  all  three  specimens. 

Fusinus  aff.  F.  ucalius  Vokes,  1939 

Figure  8 k 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  371. 

Remarks.  Only  a single  specimen  was  found  in  the  Llajas 
Formation.  It  is  unabraded  and  nearly  complete,  lacking  only 
the  protoconch.  This  specimen  agrees  with  the  description 
of  Fusinus  ucalius  Vokes  (1939:137)  in  having  swollen  col- 
labral  costae,  strong  spiral  cords,  and  a long  straight  anterior 
canal.  It  differs  in  that  the  collabral  costae  are  not  obsolete 
or  even  subobsolete  on  the  posterior  parts  of  the  spire  whorls. 
These  collabral  costae  on  the  Llajas  specimen  extend  from 
suture  to  suture.  According  to  Vokes  ( 1 939),  F.  voetus  Hanna 
(1927:315,  pi.  51,  figs.  1,  2,  4)  and  F.  meganosensis  Clark 
and  Woodford  (1927: 1 1 1,  pi.  19,  fig.  1 3)  differ  from  F.  ucali- 
us in  the  same  respect.  The  Llajas  specimen  differs  from  F. 
voetus  by  having  a less  elongate  aperture,  more  swollen  col- 
labral costae,  more  angulate  spire  whorls,  and  only  eight 
(rather  than  15)  collabral  costae  on  the  body  whorl.  The 
Llajas  specimen  differs  from  F.  meganosensis  by  having  much 
stronger  collabral  costae  and  spiral  cords.  In  addition,  on  the 


body  whorl  of  F.  meganosensis,  the  collabral  costae  tend  to 
become  obsolete  posteriorly. 

Superfamily  Volutacea 
Family  Olividae  Latreille,  1825 
Genus  Pseudoliva  Swainson,  1 840 

Type  Species.  By  original  designation,  Buccinum  plumbea 
Chemnitz,  1780?  [=Buccinum  crassa  Gmelin,  1788?]. 

Pseudoliva  lineata  Gabb,  1864 

Figure  81 

Pseudoliva  lineata  Gabb,  1864:99,  pi.  18,  fig.  52.  Stewart, 
1927:400,  pi.  28,  fig.  14a.  Vokes,  1939:139,  pi.  18,  fig.  23. 

Primary  Type  Material.  ANSP  lectotype  4200,  Tejon  For- 
mation s.l.,  northeast  of  Martinez,  California. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  Simi  Valley  through  Martinez, 
California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  373,  484. 

Remarks.  Only  three  specimens  were  found  in  the  Llajas 
Formation.  All  show  the  typical  Pseudoliva  medial  spiral 
groove.  P.  lineata  is  distinguished  by  a complete  lack  of  nodes 
on  the  shoulder  of  the  whorl,  as  noted  by  Turner  (1938:78). 
As  mentioned  by  Vokes  (1939:139),  Llajas  Formation  spec- 
imens show  a spire  that  is  somewhat  higher  than  that  of  the 
lectotype  figured  by  Stewart  (1927,  pi.  28,  fig.  14a). 

Genus  Strepsidura  Swainson,  1 840 

Type  Species.  By  original  designation,  Strepsidura  costata 
Swainson,  1840  [=Fusus  ficu/nea  Lamarck,  1822,  =Murex 
turgida  Solander,  1766], 

Strepsidura  ficus  (Gabb,  1864) 

Figure  8m 

Whitneya  ficus  Gabb,  1864:104,  pi.  28,  fig.  216.  Dickerson, 
1915:69,  pi.  9,  figs.  5a-d. 

Strepsidura  ficus  (Gabb).  Stewart,  1927:404-405,  pi.  29,  fig. 
1 1 . Kleinpell  and  Weaver,  1 963: 1 93,  pi.  27,  figs.  1 -3.  Giv- 
ens, 1974:87,  pi.  10,  fig.  10. 

Primary  Type  Material.  ANSP  lectotype  433 1 , Tejon  For- 
mation s.l..  Fort  Tejon  area,  California. 

Molluscan  Stage  Range.  Uppermost  “Capay”  through 
“Tejon.” 

Geographic  Distribution.  Simi  Valley  through  Fort  Tejon 
area,  southern  California. 


Squires,  1983a,  spire  only,  UCLA  hypotype  59195,  CSUN  loc.  444,  x0.5.  j.  Fusinus  teglandae  Hanna,  1927,  abapertural  view,  LACMIP 
hypotype  6540,  CSUN  loc.  374,  xO.75.  k.  Fusinus  aff.  F.  ucalius  Vokes,  1939,  LACMIP  hypotype  6541,  CSUN  loc.  371,  x 1.5.  I.  Pseudoliva 
lineata  Gabb,  1864,  LACMIP  hypotype  6542,  CSUN  loc.  371,  x 1.5.  m.  Strepsidura  ficus  (Gabb,  1864),  abapertural  view,  LACMIP  hypotype 
6543,  CSUN  loc.  452,  xl.25.  n.  Ancilla  ( Spirancilla ) gabbi  Cossmann,  1899,  LACMIP  hypotype  6544,  CSUN  loc.  489,  x4.  o.  Olivella 
mathewsonii  Gabb,  1864,  LACMIP  hypotype  6545,  CSUN  loc.  467,  x2.5.  p.  Proximitral  cretacea  (Gabb,  1864),  LACMIP  hypotype  6546, 
CSUN  loc.  371,  x 2.5. 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  33 


Local  Occurrence.  Interfingering  coastal  alluvial-fan  facies 
and  shallow-marine  (transgressive)  facies:  CSUN  localities 
452,  545,  703. 

Remarks.  In  some  of  the  specimens,  collabral  costae  are 
fairly  well  developed  on  the  earlier  whorls. 

The  presence  of  S.  ficus  in  the  zone  of  interfingering  be- 
tween the  coastal  alluvial-fan  facies  and  the  shallow-marine 
(transgressive)  facies  of  the  Llajas  Formation  extends  the 
molluscan  stage  range  of  this  species  into  the  uppermost 
“Capay.”  Previously,  the  lower  range  limit  had  been  reported 
as  the  “Dontengine  Stage”  (Givens,  1974). 

Subfamily  Olivinae  Swainson,  1840 

Genus  Ancilla  Lamarck,  1799 

Type  Species.  By  monotypy,  Ancilla  cinnamonea  La- 
marck, 1801. 

Ancilla  ( Spirancilla ) gabbi  Cossmann,  1899 
Figure  8n 

Ancillaria  elongata  Gabb,  1864:100,  pi.  18,  fig.  54.  Hanna, 
1927:323,  pi.  53,  figs.  9-13.  Stewart,  1927:411. 

Not  Ancillaria  elongata  Gray,  1847:357,  pi.  1,  fig.  5. 

Ancilla  gabbi  Cossmann,  1899:60  [new  name  for  Ancillaria 
elongata  Gabb,  1864,  preoccupied].  Turner,  1938:72,  pi. 
18,  fig.  6.  Weaver,  1943:500,  pi.  95,  fig.  18. 

Ancilla  (Spirancilla)  gabbi  Cossmann.  Vokes,  1939:131,  pi. 
18,  figs.  6,  10. 

Primary  Type  Material.  Holotype  lost,  UCMP  syntypes 
12521  (two  specimens)  of  Ancillaria  elongata  Gabb  and  An- 
cilla gabbi  Cossmann,  Ardath  Shale,  near  San  Diego,  perhaps 
300  m east  of  the  summit  of  Soledad  Mountain  on  the  east 
side  of  the  canyon  in  the  bottom  of  Rose  Creek. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  San  Diego,  California  through 
northwestern  Oregon. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  371.  Shallow-marine  (regressive)  facies:  CSUN 
locality  489. 

Remarks.  Only  two  specimens  were  found  in  the  Llajas 
Formation.  Both  agree  with  Gabb’s  (1864)  observations  of 
callused  sutures. 

Genus  Olivella  Swainson,  1831 

Type  Species.  By  subsequent  designation  (Dali,  1 909),  Oli- 
vella purpurata  Swainson,  1 83 1 [=Oliva  dama  Mawe,  1 823], 

Olivella  mathewsonii  Gabb,  1864 

Figure  8o 

Olivella  mathewsonii  Gabb,  1864:100,  pi.  18,  fig.  53.  An- 
derson and  Hanna,  1925:80,  pi.  8,  fig.  19.  Stewart,  1927: 
410-411,  pi.  29,  fig.  13.  Weaver,  1943:500-501,  pi.  103, 
fig.  7.  Givens,  1974:87.  Smith,  1975:469,  table  1.  Squires, 
1977:table  1. 

Primary  Type  Material.  ANSP  lectotype  4202,  Tejon  For- 
mation s.L,  Martinez,  California. 


Molluscan  Stage  Range.  Lower  “Martinez”  through  “Te- 
jon.” 

Geographic  Distribution.  Simi  Valley,  California  through 
northwestern  Washington. 

Local  Occurrence.  Interfingering  coastal  alluvial-fan  facies 
and  shallow-marine  (transgressive)  facies:  CSUN  locality 
469a.  Shallow-marine  (transgressive)  facies:  CSUN  localities 
467,  484,  498. 

Remarks.  Two  specimens  were  found  at  locality  469a.  At 
each  of  the  other  localities,  only  one  specimen  was  found. 
The  specimens  agree  with  Gabb’s  (1864)  observations  of 
three  columellar  plicae  at  the  anterior  end  of  the  shell.  This 
taxon  may  be  the  same  as  O.  m.  umpquaensis  Turner,  1938: 
72,  pi.  18,  figs.  13-14. 

Family  Mitridae  Swainson,  1831 
Subfamily  Vexillinae  Thiele,  1929 
Genus  Proximitra  Finlay,  1927 

Type  Species.  By  original  designation,  Vexillum  rutido- 
lomum  Suter,  1913. 

Proximitra ? cretacea  (Gabb,  1 864) 

Figure  8p 

Mitra  cretacea  Gabb,  1864:103,  pi.  28,  fig.  215.  Stewart, 
1927:406,  pi.  27,  figs.  9-10. 

Uromitra  (?)  cretacea  (Gabb).  Vokes,  1939:134-135,  pi.  18, 
fig.  19. 

Proximitra ? cretacea  (Gabb).  Givens,  1974:87.  Squires,  1977: 
table  1. 

Primary  Type  Material.  ANSP  holotype  4302,  Tejon  For- 
mation s.L,  Martinez,  California. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  Simi  Valley  through  central  Cal- 
ifornia. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  371. 

Remarks.  This  taxon  was  found  at  only  one  locality  in  the 
Llajas  Formation,  but  46  specimens  were  collected. 

Family  Tudiclidae  Finlay  and  Marwick,  1937 
Genus  Pseudoperissolax  Clark,  1918 

Type  Species.  By  original  designation.  Busy  coni  blakei 
Conrad,  1855. 

Pseudoperissolax  blakei  praeblakei  Vokes,  1939 

Figure  9a 

Not  Busycoril  blakei  Conrad,  1855:1  1;  1857:332,  pi.  2,  fig. 
13. 

Perissolax  blakei  (Conrad).  Gabb,  1864:92  (in  part),  pi.  21, 
fig.  110. 

Pseudoperissolax  blakei  (Conrad)  (subsp.?).  Stewart,  1927: 
429-430,  pi.  28,  fig.  1. 

Pseudoperissolax  blakei  praeblakei  Vokes,  1939:145-146,  pi. 


34  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


19,  figs.  14,  22.  Givens,  1974:88,  pi.  10,  figs.  15-16.  Smith, 
197 5:pl.  2,  fig.  16. 

Primary  Type  Material.  UCMP  holotype  15799,  UCMP 
paratype  15800,  Arroyo  Hondo  Formation,  UCMP  locality 
1817. 

Molluscan  Stage  Range.  Lower  “Martinez”  through  “Do- 
mengine.” 

Geographic  Distribution.  San  Diego?,  Simi  Valley  through 
central  California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  374. 

Remarks.  Only  two  specimens  were  found  in  the  Llajas 
Formation.  They  are  mostly  internal  molds  with  shell  present 
only  in  the  spire  area.  The  long  anterior  canal  is  missing  in 
the  figured  specimen  (Fig.  9a). 

A discussion  of  the  taxonomic  reasons  for  the  use  of  the 
family  Tudiclidae  in  place  of  the  normally  used  family  Vasi- 
dae  is  given  in  Zinsmeister  (1983b). 

Family  Harpidae  Bronn,  1 849 

Genus  Eocithara  P.  Fischer,  1883 

Type  Species.  By  monotypy,  Harpa  mutica  Lamarck,  1 803. 

Eocithara  mutica  californiensis  (Vokes,  1937) 

Figure  9b 

Harpa  ( Eocithara ) mutica  n.  sp.  Clark  and  Vokes,  1936:pl. 
2,  fig.  5. 

Harpa  (Eocithara)  mutica  californiensis  Vokes,  1937:1  1,  pi. 
2,  figs.  2,  4,  6,  8. 

Eocithara  mutica  californiensis  (Vokes).  Rehder,  1973:225, 
pi.  193. 

Primary  Type  Material.  UCMP  holotype  30438,  UCMP 
paratypes  30439-30449,  Llajas  Formation,  UCMP  locality 
3296. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  Simi  Valley,  California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  373,  374. 

Remarks.  Four  specimens  were  found  in  the  Llajas  For- 
mation. Preservation  is  generally  excellent  with  the  thin  col- 
labral  ridges  present. 

Family  Volutidae  Rafinesque,  1815 
Subfamily  Volutinae  Rafinesque,  1815 
Genus  Cryptochorda  Morch,  1858 

Type  Species.  By  monotypy,  Buccinum  stromboides 
Herrmannsen. 

Subgenus  Cryptochorda  s.s. 

Cryptochorda  (Cryptochorda)  calif ornica 
(Cooper,  1894) 

Figure  9c 

Ancilla  ( Oliverato ) californica  Cooper,  1894:43,  pi.  1,  figs. 
6-11.  Dickerson,  1913:264;  1914:115,  pi.  12,  figs.  4a-b. 


Oliverato  californica  Cooper.  Dickerson,  1913:286-287,  pi. 
1 3,  figs.  4a-b. 

Caricella  stormsiana  Dickerson,  1913:287,  pi.  13,  figs. 
3a-b. 

Cryptochorda  californica  (Cooper).  Clark,  1929:pl.  4,  figs.  6, 
16;  pi.  9,  figs.  5-6.  Clark  and  Vokes,  1936:874,  pi.  1,  fig. 
5.  Turner,  1938:72,  pi.  18,  figs.  11,  15.  Vokes,  1939:139- 
140.  Weaver,  1943:499,  pi.  95,  figs.  19,  23. 

Primary  Type  Material.  CAS  syntypes  8a-d  (four  speci- 
mens), Capay  Formation,  Marysville  Buttes,  Sutter  County, 
California. 

Molluscan  Stage  Range.  “Capay”  through  “Domengine.” 
Geographic  Distribution.  Simi  Valley,  California  through 
southwestern  Oregon. 

Local  Occurrence.  Interfingering  coastal  alluvial-fan  facies 
and  shallow-marine  (transgressive)  facies:  CSUN  locality  703. 

Remarks.  Only  two  specimens  were  found  in  the  Llajas 
Formation. 

According  to  Clark  (1921  :table  1 ; 1 929),  C.  (C.)  californica 
[=Ancilla  ( Oliverato ) californica ] occurs  in  “Meganos”-age 
strata.  Such  a report,  however,  cannot  be  substantiated  at 
this  time  because  of  his  lack  of  locality  information  and 
because  his  rock  unit  names  are  not  clearly  defined. 

Genus  Lyria  Gray,  1847 

Type  Species.  By  original  designation,  Voluta  nucleus  La- 
marck, 1811. 

Lyria  andersoni  Waring,  1917 

Figure  9d 

Cance/laria  irelaniana  Cooper,  1 894:42,  pi.  1,  fig.  5.  Arnold, 
1910:52,  pi.  4,  fig.  22.  [Misidentification.] 

Lyria  andersoni  Waring,  1917:97,  pi.  15,  fig.  12. 

Lyria  andersoni  Waring.  Clark,  1929:pl.  9,  figs.  7-8.  Clark 
and  Vokes,  1936:876,  pi.  1,  fig.  17.  Turner,  1938:73,  pi. 
1 8,  fig.  5.  Vokes,  1 939: 1 36,  pi.  1 8,  figs.  22,  24.  Hanna  and 
Hertlein,  1943:170,  fig.  62-21. 

Primary  Type  Material.  SU  holotype  195,  SU  paratype 
196,  Llajas  Formation,  SU  locality  2696. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  Simi  Valley,  California  through 
southwestern  Oregon. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  372,  439,  455,  466,  484,  498,  499,  507. 

Remarks.  This  taxon  helps  to  characterize  the  lower  and 
middle  parts  of  the  shallow-marine  (transgressive)  facies.  It 
is  most  common  in  the  middle  part.  Preservation  is  generally 
good  to  excellent. 

Subfamily  Fulgorariinae 
Pilsbry  and  Olsson,  1954 

Genus  Lyrischapa  Aldrich,  1911 

Type  Species.  By  monotypy,  Lyrischapa  harrisi  Aldrich, 
1911. 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  35 


Lyrischapa  lajollaensis  (Hanna,  1927) 

Figure  9e 

Pejonia  lajollaensis  Hanna,  1927:320,  pi.  52,  figs.  1-2. 
Volutospira  ( Pejonia ) lajollaensis  (Hanna).  Clark,  1929,  pi. 

9,  figs.  11-12. 

Volutocristata  lajollaensis  (Hanna).  Gardner  and  Bowles, 
1934:246,  fig.  13.  Givens,  1974:88. 

Lyrischapa  lajollaensis  (Hanna).  Givens,  1979:124-126,  pi. 
3,  figs.  1-2;  pi.  4,  figs.  1-3.  Givens  and  Kennedy,  1979: 
table  1. 

Primary  Type  Material.  Holotype  lost,  UCMP  neotype 
14634,  Ardath  Shale,  UCMP  locality  5062. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  San  Diego  through  Pine  Moun- 
tain area,  southern  California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  373,  374,  444,  445,  510,  543,  704. 

Remarks.  This  taxon  is  most  common  in  the  “Stewart  bed” 
at  locality  374,  where  some  rather  large  specimens  (47  mm 
width)  occur. 

Hypotype  specimens  figured  by  Givens  (1979)  are  from 
the  Llajas  Formation. 

Family  Cancellariidae 
Forbes  and  Hanley,  1853 

Genus  Bonellitia  Jousseaume,  1887 

Type  Species.  By  original  designation,  Cance/laria  bonellii 
Bellardi,  1872? 

Subgenus  Admetula  Cossmann,  1889 

Type  Species.  By  original  designation,  Cancellaria  evulsa 
Solander,  1776. 

Bonellitia  ( Admetula ) paucivaricata 
(Gabb,  1864) 

Figure  9f 

Tritonium  paucivaricatum  Gabb,  1864:95,  pi.  28,  figs.  209, 
209a. 

Cancellaria  stantoni  Dickerson,  1913:282,  pi.  12,  figs.  2a-b. 
Ad  mete  (Bonellitia)  stantoni  (Dickerson).  Weaver  and  Palm- 
er, 1922:40-42,  pi.  11,  figs.  1,  5. 

Cancellaria  paucivaricata  (Gabb).  Anderson  and  Hanna, 
1925:81,  pi.  8,  figs.  3-4. 

Bonellitia  (Admetula)  paucivaricata  (Gabb).  Stewart,  1927: 
413,  pi.  29,  fig.  5.  Turner,  1938:71,  pi.  15,  figs.  12-13. 
Weaver,  1943:508,  pi.  96,  figs.  10,  12,  13;  pi.  103,  figs.  9, 

10,  17.  Givens,  1974:89.  Squires,  1977:table  1. 

Primary  Type  Material.  ANSP  lectotype  4 1 94,  Tejon  For- 
mation, Tejon  Pass  area,  California. 

Molluscan  Stage  Range.  “Meganos”?,  “Capay”  through 
“Tejon.” 

Geographic  Distribution.  Simi  Valley,  California  through 
southwestern  Washington. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  371. 


Remarks.  Only  seven  specimens  were  found  in  the  Llajas 
Formation.  According  to  Clark  (1921:table  1;  1926:1  15),  B. 
(A.)  paucivaricata  [= Cancellaria  stantoni  and  Admeta  stan- 
toni] occurs  in  “Meganos”-age  strata.  Such  a report,  however, 
cannot  be  substantiated  at  this  time  because  of  his  lack  of 
locality  information  and  because  his  rock  unit  names  are  not 
clearly  defined.  Clark  and  Woodford  (1927),  however,  re- 
ported a questionable  occurrence  of  B.  (A.)  paucivaricata 
[=Admete  (Bonellitia)  cf.  stantoni ] from  localities  of  late  Pa- 
leocene/early  Eocene  age  strata. 

Superfamily  Conacea 
Family  Turridae  Swainson,  1840 
Subfamily  Turriculinae  Powell,  1942 
Genus  Pleurofusia  de  Gregorio,  1890 

Type  Species.  By  original  designation,  Pleurotoma  (Pleu- 
rofusia) longirostropis  de  Gregorio,  1890. 

Pleurofusia  fresnoensis  (Arnold,  1910) 

Figure  9g 

Pleurotoma  fresnoensis  Arnold,  1910:53,  pi.  4,  fig.  23. 
Surcu/a  clarki  Dickerson,  1913:278,  pi.  1 1,  fig.  3. 
Pleurofusia  fresnoensis  (Arnold).  Vokes,  1939:1  17-1  18,  pi. 

17,  figs.  15-16.  Givens,  1974:90,  pi.  11,  fig.  9.  Givens  and 

Kennedy,  1979:95,  tables  1,  3. 

Primary  Type  Material.  USNM  holotype  165631,  Do- 
mengine Formation,  USGS  locality  4619. 

Molluscan  Stage  Range.  “Capay”  through  “Transition.” 

Geographic  Distribution.  San  Diego  through  central  Cal- 
ifornia. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  538,  539. 

Remarks.  Specimens  are  most  common  at  locality  371. 

According  to  Clark  (1921  :table  1 ; 1 926: 1 1 6),  P.  fresnoensis 
[=Turris  (Surcu/a)  clarki  and  Surcula  clarki]  occurs  in  “Me- 
ganos”-age  strata.  Such  a report,  however,  cannot  be  sub- 
stantiated at  this  time  because  of  his  lack  of  locality  infor- 
mation and  because  his  rock  unit  names  are  not  clearly  defined. 

Smith  (1975:table  1)  reported  Pleurofusia  sp.  aff.  P.  fres- 
noensis from  “Martinez”-age  strata  of  central  California. 

Genus  Fusiturricula  Woodring,  1928 

Type  Species.  By  original  designation,  Turris  (Surcula) 
fusinella. 

Subgenus  Crenaturricula  Vokes,  1939 

Type  Species.  By  original  designation,  Surcula  crenato- 
spira  Cooper,  1894. 

Fusiturricula  ( Crenaturricula ) crenatospira 
(Cooper,  1894) 

Figure  9h 

Surcula  crenatospira  Cooper,  1894:39,  pi.  1,  figs.  2-4.  Dick- 
erson, 1913:278,  pi.  11,  fig.  4. 


36  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Fusiturricula  ( Crenaturricula ) crenatospira  (Cooper).  Vokes, 
1939:1  14-115,  pi.  17,  figs.  4-5. 

Primary  Type  Material.  CAS  syntypes  9a-b  (two  speci- 
mens), Capay  Formation,  UCMP  locality  1853. 

Molluscan  Stage  Range.  “Capay”  through  “Domengine.” 
Geographic  Distribution.  Simi  Valley  through  Marysville 
Buttes,  California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  374. 

Remarks.  Only  two  specimens  were  found,  and  these  are 
from  the  “Stewart  bed.”  According  to  Vokes  (1939:15)  this 
taxon  is  confined  to  and  is  diagnostic  of  the  “Capay  Stage.” 
The  presence  of  it  in  the  “Stewart  bed”  extends  its  provincial 
range  into  the  “Domengine  Stage.” 

According  to  Clark  (1921:table  1;  1929),  F.  (C.)  crenato- 
spira [=Turris  ( Surcula ) crenatospira  and  Turricula  crenato- 
spira] occurs  in  “Meganos”-age  strata.  Such  a report,  how- 
ever, cannot  be  substantiated  at  this  time  because  of  his  lack 
of  locality  information  and  because  his  rock  unit  names  are 
not  clearly  defined. 

Fusiturricula  ( Crenaturricula ) crenatospira 
domenginica  Vokes,  1939 

Figure  9i 

Fusiturricula  { Crenaturricula ) crenatospira  domenginica 
Vokes,  1939:1  15-1  16,  pi.  17,  figs.  6-7. 

Primary  Type  Material.  UCMP  holotype  15768,  Llajas 
Formation,  UCMP  locality  7002;  UCMP  paratype  15769, 
Llajas  Formation,  UCMP  locality  3304. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  Simi  Valley  through  central  Cal- 
ifornia. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  374,  444,  445,  498,  548. 

Remarks.  At  locality  374,  this  taxon  occurs  along  with  F. 
(C.)  crenatospira,  which  is  characterized  by  the  presence  of 
collabral  sculpturing  below  the  nodes  on  the  spire  and  body 
whorls.  On  F.  ( C .)  crenatospira  domenginica,  the  collabral 
sculpturing  is  obsolete.  Contrary  to  Vokes’  (1939:1  15-116) 
comments,  the  two  taxa  have  approximately  the  same  num- 
ber of  nodes  on  the  body  whorl. 

Genus  Surculites  Conrad,  1865 

Type  Species.  By  monotypy,  Surcula  (Surculites)  annosus 
Conrad,  1865. 

Surculites  mathewsonii  (Gabb,  1864) 

Figure  9j 

Fusus  mathewsonii  Gabb,  1 864:83,  pi.  1 8,  fig.  33.  Dickerson, 
1 9 1 4:pl.  16,  fig.  2. 

Bela  clathra  Gabb,  1869:152,  pi.  26,  fig.  31. 

IPieurotoma  decipiens  Cooper,  1894:40,  pi.  2,  fig.  32. 
Potamidesl  davisiana  Cooper,  1894:44,  pi.  1,  fig.  13. 
Surcula  davisiana  (Cooper).  Dickerson,  1 9 1 3:279,  pi.  12,  figs. 
6a-b. 


Surcula  (?)  sp.  Waring,  1917 :pl.  15,  fig.  16. 

Surcula  decipiens  (Cooper).  Hanna,  1927:324,  pi.  54,  figs. 

6,  8. 

Surculites  mathewsonii  (Gabb).  Stewart,  1927:420-421,  pi. 

26,  figs.  12-14.  Clark,  1929:pl.  9,  figs.  3-4.  Turner,  1938: 
69-70,  pi.  17,  figs.  6,  10.  Vokes,  1939:123,  pi.  17,  figs.  8, 
19.  Weaver,  1943:526,  pi.  97,  figs.  24,  29;  pi.  98,  figs.  1, 
5;  1953:29.  Givens,  1974:90,  pi.  11,  figs.  5,  7.  Zinsmeister, 
1974:164,  pi.  17,  fig.  6;  1983a:table  1.  Squires,  1977:table 
1.  Givens  and  Kennedy,  1979:87,  tables  1,  3. 

“ Surculites ” mathewsonii  (Gabb).  Smith,  1975:pl.  2,  fig.  15. 

Primary  Type  Material.  ANSP  lectotype  4180,  Tejon  For- 
mation s.l.,  near  Martinez,  California. 

Molluscan  Stage  Range.  Lower  “Martinez”  through 
“Transition.” 

Geographic  Distribution,  San  Diego,  California  through 
southwestern  Oregon. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  372,  374,  444,  445,  447,  479,  498. 

Remarks.  Specimens  are  fairly  common  at  locality  37 1 but 
are  more  common  and  larger  at  locality  374  of  the  “Stewart 
bed.”  At  locality  374,  the  specimens  are  internal  molds.  Spec- 
imens at  locality  371  are  strongly  angulated,  which  is  in 
keeping  with  one  of  the  main  characteristics  of  this  species 
as  discussed  by  Vokes  (1939:123). 

Genus  Domenginella  Vokes,  1939 

Type  Species.  By  original  designation,  Tunis  claytonensis 
Gabb,  1864. 

Domenginella  claytonensis  (Gabb,  1864) 

Figure  9k 

Turris  claytonensis  Gabb,  1864:92,  pi.  18,  fig.  46. 

Surcula  gesteri  Dickerson,  1916:499,  pi.  42,  fig.  4. 
Scobinella  claytonensis  ( Gabb).  Stewart,  1927:417-418,  pi. 

27,  figs.  11-12. 

Domenginella  claytonensis  (Gabb).  Vokes,  1939:122-123,  pi. 
17,  figs.  18,  20. 

Primary  Type  Material.  ANSP  lectotype  4 1 90,  Tejon  For- 
mation s.l.,  near  Martinez,  California. 

Molluscan  Stage  Range.  Uppermost  “Capay”  through 
“Domengine.” 

Geographic  Distribution.  Simi  Valley  through  central  Cal- 
ifornia. 

Local  Occurrence.  Interfingering  coastal  alluvial-fan  facies 
and  shallow-marine  (transgressive)  facies:  CSUN  locality 
469a.  Shallow-marine  (transgressive)  facies:  CSUN  localities 
371,  372,  455,  466,  484,  486,  498,  517,  538,  539,  548. 

Remarks.  This  taxon  is  most  common  at  locality  371, 
where  most  of  the  specimens  are  well-preserved  juveniles. 

The  presence  of  D.  claytonensis  in  the  zone  of  interfingering 
between  the  coastal  alluvial-fan  facies  and  the  shallow-ma- 
rine (transgressive)  facies  of  the  Llajas  Formation  extends 
the  molluscan  stage  range  of  this  taxon  into  the  uppermost 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  37 


Figure  9.  Llajas  Formation  gastropods  (continued).  Unless  otherwise  indicated,  views  are  apertural.  a.  Pseudoperissolax  blakei  praeblakei 
Vokes,  1939,  internal  mold,  side  view,  LACMIP  hypotype  6547,  CSUN  loc.  374,  xl.5.  b.  Eocithara  mutica  californiensis  (Vokes,  1937), 
abapertural  view,  LACMIP  hypotype  6548,  CSUN  loc.  373,  xl.5.  c.  Cryptochorda  (Cryptochorda)  californica  (Cooper,  1894),  abapertural 
view,  LACMIP  hypotype  6549,  CSUN  loc.  703,  x 1.25.  d.  Lyria  andersoni  Waring,  1917,  LACMIP  hypotype  6550,  CSUN  loc.  498,  x2.  e. 
Lyrischapa  lajollaensis  (Hanna,  1927),  LACMIP  hypotype  6551,  CSUN  loc.  373,  xl.  f.  Bonellita  (Admetula)  paucivariata  (Gabb,  1864), 
LACMIP  hypotype  6552,  CSUN  loc.  374,  x4.5.  g.  Pleurofusia  fresnoensis  (Arnold,  1910),  LACMIP  hypotype  6553,  CSUN  loc.  371,  x3.  h. 
Fusiturricula  (Crenaturricula)  crenatospira  (Cooper,  1894),  abapertural  view,  UCLA  hypotype  59278,  CSUN  loc.  374,  x 1.5.  i.  Fusiturricula 
( Crenaturricula ) crenatospira  domenginica  Vokes,  1939,  abapertural  view,  LACMIP  hypotype  6554,  CSUN  loc.  371,  xl.  j.  Surculites  ma- 


38  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


“Capay.”  Previously,  the  lower  range  limit  had  been  reported 
as  the  “Domengine  Stage”  (Vokes,  1939). 

Genus  Exilia  Conrad,  1860 

Type  Species.  By  monotypy,  Exilia  pergracilis  Conrad, 
1860. 

Exilia  llajasensis  Bentson,  1940 

Figure  91 

Exilia  llajasensis  Bentson,  1940:212,  pi.  1,  fig.  28;  pi.  2,  fig. 
16. 

Primary  Type  Material.  UCMP  holotype  12144,  Llajas 
Formation,  UCMP  locality  A-3042;  UCMP  paratype  12140, 
Llajas  Formation,  UCMP  locality  3296. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  Simi  Valley,  California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  486,  539. 

Remarks.  Only  two  specimens  were  found  in  the  Llajas 
Formation.  In  the  primary  type  specimens,  the  anterior  canal 
is  missing.  As  can  be  seen  in  Figure  91,  the  long  anterior  canal 
is  covered  by  10  spiral  ribs. 

Family  Conidae  Rafinesque,  1815 
Genus  Conus  Linne,  1758 

Type  Species.  By  subsequent  designation  (Children,  1 823), 
Conus  marmoreus  Linne,  1758. 

Conus  caleocius  Yokes,  1939 

Figure  9m 

Conus  caleocius  Vokes,  1939:127-129,  pi.  18,  figs.  1,  7. 

Primary  Type  Material.  UCMP  holotype  15785,  Llajas 
Formation,  UCMP  locality  3310. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  Simi  Valley  through  central  Cal- 
ifornia. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  498. 

Remarks.  Four  specimens  were  found.  Three  of  these  are 
from  locality  371. 

Conus  hornii  umpquaensis  Turner,  1938 

Figure  9n 

Conus  hornii  umpquaensis  Turner,  1938:69,  pi.  15,  figs.  1- 
2.  Vokes,  1939:127,  pi.  18,  figs.  2-3.  Stewart,  1946:pl.  1 1, 


fig.  6.  Weaver,  1943:510-51 1,  pi.  96,  fig.  18.  Givens  and 

Kennedy,  1979:87,  tables  1,  3. 

Primary  Type  Material.  UCMP  holotype  33656,  “upper 
Umpqua”  Formation,  UO  locality  144. 

Molluscan  Stage  Range.  “Domengine”  through  “Transi- 
tion.” 

Geographic  Distribution.  San  Diego,  California  through 
southwestern  Oregon. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  37 1 , 373,  374,  449,  455,  484,  486,  49 1 , 498, 
538,  539. 

Remarks.  This  taxon  characterizes  the  lower  half  of  the 
shallow-marine  (transgressive)  facies.  It  is  especially  abun- 
dant at  locality  371,  where  it  forms  a growth  series. 

Family  Terebridae  H.  and  A.  Adams,  1854 
Genus  Terebra  Bruguiere,  1789 

Type  Species.  By  subsequent  designation  (Lamarck,  1 799), 
Buccinum  subulata  Linne,  1758. 

Terebra  californica  Gabb,  1869 

Figure  9o 

Terebra  californica  Gabb,  1869:162,  pi.  27,  fig.  41. 

Terebra  californica  Gabb.  Anderson  and  Hanna,  1925:82, 

pi.  8,  fig.  18.  Stewart,  1927:424,  pi.  26,  fig.  5.  Vokes,  1939: 

113,  pi.  16,  fig.  38.  Givens,  1974:92. 

Primary  Type  Material.  ANSP  holotype  4209,  Tejon  For- 
mation s.l.,  Martinez,  California. 

Molluscan  Stage  Range.  “Domengine”  through  “Tejon.” 

Geographic  Distribution.  Simi  Valley  through  central  Cal- 
ifornia. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  467. 

Remarks.  Only  two  specimens  were  found.  They  are  very 
small,  with  heights  of  3 mm. 

Subclass  Euthyneura 
Order  Cephalaspidea 

Superfamily  Cylichnacea  A.  Adams,  1850 
Family  Cylichnidae  A.  Adams,  1850 
Genus  Cy/ichnina  Monterosato,  1884 

Type  Species.  By  original  designation.  Bulla  umbilicata 
Montagu,  1803. 


thewsonii  (Gabb,  1864),  abapertural  view,  LACMIP  hypotype  6555,  CSUN  loc.  371,  x2.5.  k.  Domenginella  claytonensis  (Gabb,  1864),  side 
view,  LACMIP  hypotype  6556,  CSUN  loc.  371,  x2.5.  1-  Exilia  llajasensis  Bentson,  1940,  LACMIP  hypotype  6557,  CSUN  loc.  539,  x2.5. 
m.  Conus  caleocius  V okes,  1939,  LACMIP  hypotype  6558,  CSUN  loc.  498,  x 2.  n.  Conus  hornii  umpquaensis  Turner,  1938,  LACMIP  hypotype 
6559,  CSUN  loc.  371,  x2.5.  o.  Terebra  californica  Gabb,  1869,  LACMIP  hypotype  6560,  CSUN  loc.  467,  x 8.  p.  Cylichnina  tantilla  (Anderson 
and  Hanna,  1925),  LACMIP  hypotype  6561,  CSUN  loc.  371,  x2.5.  q.  Scaphander  (Mirascapha)  costatus  (Gabb,  1864),  abapertural  view, 
LACMIP  hypotype  6562,  CSUN  loc.  493,  x 1.5.  r.  Megistostoma  gabbianum  (Stoliczka,  1868),  internal  mold,  abapertural  view,  LACMIP 
hypotype  6553,  CSUN  loc.  458,  x 1.5. 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  39 


Cylichnina  tantilla 
(Anderson  and  Hanna,  1925) 

Figure  9p 

Cylichnella  tantilla  Anderson  and  Hanna,  1925:140,  pi.  7, 
figs.  4,  8-9. 

Cylichnina  tantilla  (Anderson  and  Hanna).  Stewart,  1927: 
439-441,  pi.  27,  figs.  2-4.  Turner,  1938:67-68,  pi.  20,  figs. 
9-10.  Vokes,  1939:110,  pi.  16,  figs.  28,  33,  39.  Weaver, 
1943:548-549,  pi.  100,  figs.  10-12,  1 4-1 5.  Stewart,  1946: 
pi.  11,  fig.  11.  Givens,  1974:93.  Squires,  1977:table  1; 
1983b,  fig.  9a. 

Primary  Type  Material.  CAS  holotype  958,  CAS  paratypes 
959  and  960,  Tejon  Formation,  CAS  locality  711. 

Molluscan  Stage  Range.  “Domengine”  through  “Tejon.” 
Geographic  Distribution.  Simi  Valley,  California  through 
western  Washington. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  372,  373,  374,  455,  460,  466,  467, 
469e,  469f,  484,  486,  491,  492,  498,  499,  507,  508,  517, 
537,  538,  540,  543.  Shallow-marine  (regressive)  facies:  CSUN 
locality  475. 

Remarks.  This  easily  recognized  taxon  is  especially  abun- 
dant at  localities  371,  373,  and  455.  At  these  localities  it 
occurs  as  growth  series. 

Genus  Scaphander  Montfort,  1810 

Type  Species.  By  original  designation.  Bulla  lignaria  Linne, 
1767. 

Subgenus  Mirascapha  Stewart,  1927 

Type  Species.  By  original  designation,  Cylichna  costata 
Gabb,  1864. 

Scaphander  ( Mirascapha ) costatus 
(Gabb,  1864) 

Figure  9q 

Cylichna  costata  Gabb,  1864:143-144,  pi.  21,  fig.  107.  Ar- 
nold, 1 907 :pl.  39,  fig.  10.  Zinsmeister,  1974:170-171,  pi. 
12,  figs.  20-22. 

Scaphander  costata  (Gabb).  Hanna,  1927:329,  pi.  57,  figs.  2, 
3,  5. 

Scaphander  (Mirascapha)  costatus  (Gabb).  Stewart,  1927: 
437-438,  pi.  27,  fig.  5.  Turner,  1938:67,  pi.  17,  fig.  16. 
Vokes,  1939:109,  pi.  16,  figs.  29,  35.  Givens,  1974:93-94. 
Scaphander  costatus  (Gabb).  Weaver,  1943:545,  pi.  100,  fig. 
2;  pi.  103,  fig.  21.  Weaver,  1953:29.  Givens  and  Kennedy, 
1979:88,  table  3. 

Primary  Type  Material.  ANSP  lectotype  4338,  Tejon  For- 
mation s.l.,  near  Martinez,  California. 

Molluscan  Stage  Range.  “Martinez”  through  “Transi- 
tion.” 

Geographic  Distribution.  San  Diego,  California  through 
western  Washington. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 


CSUN  locality  493.  Shallow-marine  (regressive)  facies:  CSUN 
locality  475. 

Remarks.  Only  a few  specimens  were  found  in  the  Llajas 
Formation. 

Family  Philinidae  Gray,  1850 

Genus  Megistostoma  Gabb,  1864 

Type  Species.  By  monotypy,  Megistostoma  striata  Gabb, 
1864. 

Megistostoma  gabbianum  (Stoliczka,  1868) 

Figure  9r 

Megistostoma  striata  Gabb,  1864:144,  pi.  21,  figs.  108a-b. 
Not  Bullaea  striata  Deshayes,  1824:37,  pi.  5,  figs.  1-3. 
Bullaea  gabbiana  Stoliczka,  1868:434  [new  name  for  Meg- 
istostoma striata  Gabb,  1864,  preoccupied]. 

Philine (Megistostoma) gabbiC ossmann,  1895:127  [new  name 
for  Megistostoma  striata  Gabb,  1864,  preoccupied], 
Megistostoma  gabbianum  (Stoliczka).  Stewart,  1 927:44 1-442, 
pi.  26,  figs.  1-2.  Vokes,  1939:112,  pi.  17,  figs.  1-3.  Givens 
and  Kennedy,  1979:88,  table  3. 

Megistostoma  caminoensis  Hanna,  1927:330,  pi.  57,  figs.  9- 
10.  Turner,  1938:68,  pi.  20,  fig.  15.  Weaver,  1943:541,  pi. 
99,  fig.  24. 

Primary  Type  Material.  ANSP  holotype  4216  of  Megis- 
tostoma striata  Gabb,  Bullaea  gabbiana  Stoliczka,  and  Phi- 
line  (Megistostoma)  gabbi  Cossmann,  Tejon  Formation  s.l., 
near  Martinez,  California. 

Molluscan  Stage  Range.  “Domengine”  through  “Transi- 
tion.” 

Geographic  Distribution.  San  Diego,  California  through 
southwestern  Oregon. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  374, 458.  Shallow-marine  (regressive)  facies: 
CSUN  locality  462. 

Remarks.  Only  a few  specimens  were  found  in  the  Llajas 
Formation,  and  they  occur  as  internal  molds. 

Class  Bivalvia 
Subclass  Palaeotaxodonta 
Order  Nuculoida 
Superfamily  Nuculacea 
Family  Nuculidae  Gray,  1824 
Genus  Acila  H.  & A.  Adams,  1858 

Type  Species.  By  subsequent  designation  (Stoliczka,  1871), 
Nucula  divaricata  Hinds,  1843. 

Subgenus  Trunc acila  Schenck,  in 
Grant  and  Gale,  1931 

Type  Species.  By  original  designation  Nucula  castrensis 
Hinds,  1843. 


40  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Acila  ( Truncacila ) decisa  (Conrad,  1855) 

Figure  10a 

Nucula  decisa  Conrad,  1855:1  1-12;  1 8 5 7:pl.  3,  fig.  19. 

Acila  gabbiana  Dickerson,  1916:481,  pi.  36,  fig.  1.  Anderson 
and  Hanna,  1925:176,  pi.  9,  fig.  12. 

Nucula  {Acila)  stil/waterensis  Weaver  and  Palmer,  1922:6, 
pi.  8,  fig.  8. 

Acila  lajollaensis  Hanna,  1927:270,  pi.  25,  figs.  1,  3,  5,  7-8, 
12,  15. 

Acila  {Truncacila)  decisa  (Conrad).  Schenck,  1 936:53-56,  pi. 
3,  figs.  1-9,  11-15;  pi.  4,  figs.  1-2;  text  figure  7 (22,  23, 
25).  Turner,  1938:41-42,  pi.  5,  figs.  2-3.  Vokes,  1939:41, 
pi.  1,  figs.  7-8.  Weaver,  1943:22-23,  pi.  6,  figs.  1,  4,  8;  pi. 
7,  figs.  8-9.  Givens,  1974:38,  pi.  1,  fig.  1.  Zinsmeister, 
1974:67-68,  pi.  6,  fig.  3.  Squires,  1977:table  1.  Moore, 
1983:A10,  pi.  1,  fig.  14.  Zinsmeister,  1983a:table  1. 

Primary  Type  Material.  UCMP  neotype  31132,  Ardath 
Shale,  UCMP  locality  5062. 

Molluscan  Stage  Range.  “Martinez”  through  upper  Eocene 
{Turritella  schencki  delaguerrae Zone  of  Kleinpell  and  Weav- 
er, 1963). 

Geographic  Distribution.  San  Diego,  California  through 
Kamchatka,  Alaska. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  467. 

Remarks.  Specimens  are  abundant  at  locality  467.  Pres- 
ervation is  excellent.  The  specimens  are  disarticulated,  but 
the  valves  constitute  a growth  series. 

Superfamily  Nuculanacea 
Family  Nuculanidae  H.  and  A.  Adams,  1858 
Genus  Nuculana  Link,  1 807 

Type  Species.  By  original  designation.  Area  rostrata 
Chemnitz,  1784  [=Arca  pernula  Muller,  1776], 

Subgenus  Saccella  Woodring,  1925 

Type  Species.  By  original  designation.  Area  fragilis  Chem- 
nitz, 1784. 

Nuculana  ( Saccella ) gabbii  (Gabb,  1864) 

Figure  10b 

Not  Leda  protextal  Gabb,  1860:303,  pi.  48,  fig.  23. 

Leda  protextal  Gabb,  1864:199  (in  part),  pi.  26,  fig.  185. 
Nuculana  gabbii  Conrad,  1866:3  [ nomen  nudum], 

Leda  gabbii  Conrad.  Gabb,  1869:197  [new  name  for  Leda 
protextal  Gabb,  1864,  misidentified].  Stanton,  1896:1041, 
pi.  64,  fig.  8.  Waring,  1917:76,  pi.  13,  fig.  6.  Dickerson, 
191 5:pl.  1,  fig.  1;  1916,  pi.  36,  fig.  3.  Clark  and  Woodford, 
1927:85-86,  pi.  14,  fig.  2.  Clark,  1929:pl.  3,  fig.  12. 

Leda  vaderensis  Dickerson,  1915:52,  pi.  1,  fig.  3. 

Leda  vogdesi  Anderson  and  Hanna,  1925: 1 77-1 79,  pi.  2,  figs. 
8-9. 

Saccella  gabbii  (Gabb).  Stewart,  1930:55-58,  pi.  7,  fig.  3 [not 
pi.  10,  fig.  4.  =N.  (S'.)  alaeformis  fide  Zinsmeister,  1974], 
Weaver,  1953:28. 


Nuculana  {Saccella)  gabbii  (Gabb).  Vokes,  1939:41-42. 
Kleinpell  and  Weaver,  1963:195,  pi.  28,  fig.  1.  Givens, 
1974:39,  pi.  1,  fig.  3.  Moore,  1983:A16,  pi.  2,  figs.  7-8. 
Nuculana (Calorhadia)  gabbii { Gabb).  Zinsmeister,  1974:69- 
70,  pi.  6,  fig.  7;  1983a,  pi.  1,  fig.  3. 

Primary  Type  Material.  ANSP  lectotype  4476  of  Leda 
gabbii  Gabb,  1869,  Tejon  Formation  s.l.,  Martinez,  Califor- 
nia. 

Molluscan  Stage  Range.  “Martinez”  through  upper  Eocene 
( T urritella  schencki  delaguerrae  Zone  of  Kleinpell  and  Weav- 
er, 1963). 

Geographic  Distribution.  Simi  Valley,  California  through 
Kamchatka,  Alaska. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  467. 

Remarks.  Only  five  disarticulated  valves  were  found  in  the 
Llajas  Formation.  There  is  considerable  variation  in  the 
amount  of  elongation  of  the  rostrum  in  these  specimens,  with 
the  illustrated  specimen  (Fig.  10b)  showing  the  most. 

According  to  Zinsmeister  (1974),  Stewart  ( 1930)  somehow 
confused  N.  {S.)  gabbii  with  N.  {S.)  alaeformis  and  designated 
N.  (S.)  alaeformis  (pi.  10,  fig.  4)  as  the  lectotype  (ANSP  4476) 
of  N.  {S.)  gabbii. 

Subclass  Pteriomorpha 
Order  Arcoida 
Superfamily  Limnopsacea 
Family  Glycymerididae  Newton,  1922 
Subfamily  Glycymeridinae  Newton,  1922 
Genus  Glycymeris  da  Costa,  1778 

Type  Species.  By  tautonymy.  Area  orbicularis  da  Costa, 
1778  [=Arca  glycymeris  Linne,  1758], 

Subgenus  Glycymeris  s.s. 

Glycymeris  ( Glycymeris ) rosecanyonensis 
Hanna,  1927 
Figure  10c 

Glycymeris  rosecanyonensis  Hanna,  1927:273-274,  pi.  27, 
figs.  4-5,  9,  1 1.  Clark,  1929:pl.  6,  fig.  8.  Givens  and  Ken- 
nedy, 1979:tables  1,  3. 

Glycymeris  { Glycymeris ) rosecanyonensis  Hanna.  Givens, 
1974:42.  Moore,  1983:A49-A50,  pi.  10,  figs.  12-13. 
Glycymeris  { Glycymerisa ) rosecanyonensis  Hanna.  Squires, 
1977:table  1. 

Primary  Type  Material.  UCMP  holotype  30989,  Ardath 
Shale,  UCMP  locality  3990. 

Molluscan  Stage  Range.  “Domengine”  through  “Transi- 
tion.” 

Geographic  Distribution.  San  Diego  through  lower  Piru 
Creek,  southern  California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  41 


CSUN  localities  373,  467,  47 Id,  520.  Shallow-marine  (re- 
gressive) facies:  CSUN  localities  476,  489,  512a,  512b. 

Remarks.  Only  a few  specimens  of  this  small-sized  bivalve 
were  found  in  the  Llajas  Formation. 

Subgenus  Glycymerita 
Finlay  and  Marwick,  1937 

Type  Species.  By  original  designation,  Glycymeris  concava 
Marshall,  1917. 

Glycymeris  ( Glycymerita ) sagittata 
(Gabb,’  1864) 

Figure  lOd 

Axinaea  ( Limopsis ?)  sagittata  Gabb,  1864:197-198,  pi.  31, 
figs.  267,  267a. 

Glycimeris  hannibali  Dickerson,  1916:483,  pi.  36,  figs. 
8a-b. 

Glycymeris  sagittatus  (Gabb).  Dickerson,  1 9 1 6:pl.  36,  figs. 
5a-b. 

Glycymeris  sagittata  (Gabb).  Anderson  and  Hanna,  1925: 
181-182,  pi.  1,  fig.  6.  Stewart,  1930:71-73,  pi.  12,  fig.  10; 
1946:pl.  12,  fig.  3.  Vokes,  1939:45-46,  pi.  1,  figs.  18-20. 
Weaver,  1943:54-55,  pi.  9,  figs.  17-18;  pi.  11,  fig.  15. 
Kleinpell  and  Weaver,  1963:196-197,  pi.  28,  fig.  10;  pi. 
29,  figs.  1-2. 

Glycimeris  sagittatus  (Gabb).  Turner,  1938:43-44,  pi.  6,  figs. 

1-3. 

Glycymeris  ( Glycymerita ) sagittata  (Gabb).  Givens,  1974: 
42-43.  Squires,  1977:table  1.  Moore,  1 983:A54-A55,  pi. 
12,  fig.  17. 

Primary  Type  Material.  ANSP  lectotype  4422,  Tejon  For- 
mation, near  Fort  Tejon  (N  Vi  of  section  29,  T 10  N,  R 19 
W,  Kern  County,  California). 

Molluscan  Stage  Range.  “Capay”  through  “Tejon,”  Oli- 
gocene?. 

Geographic  Distribution.  Simi  Valley,  California  through 
southwestern  Washington. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  373,  374,  444,  500,  704. 

Remarks.  Specimens  are  uncommon  in  the  Llajas  For- 
mation, and  leaching  has  obscured  some  of  the  exterior  sculp- 
ture. 

Order  Mytiloida 
Superfamily  Mytilacea 
Family  Mytilidae  Rafinesque,  1815 
Genus  Brachidontes  Swainson,  1840 

Type  Species.  By  monotypy,  Modiola  sulcata  Lamarck, 
1819  (not  1 805)  [=Mytilus  citrinus  Roding,  1 798,  =Arca  mo- 
diolus Linne,  1767], 


Subgenus  Brachidontes  s.s. 

Brachidontes  ( Brachidontes ) cowlitzensis 
(Weaver  and  Palmer,  1922) 

Figure  lOe 

Modiola  ornata  Gabb,  1864:184-185,  pi.  24,  fig.  166. 

Not  Mytilus  ornatus  Orbigny,  1843:283,  pi.  342,  figs.  10-12. 
Modiolus  ornatus  Gabb.  Arnold,  1907:pl.  38,  fig.  4. 
Brachydontes  ornatus  (Gabb).  Anderson  and  Hanna,  1925: 
188,  pi.  3,  fig.  4. 

Modiolus  ( Brachydontes ) ornatus  Gabb.  Clark  and  Wood- 
ford, 1927:89,  pi.  14,  fig.  10.  Clark,  1929:pl.  3,  fig.  6. 
Modiolus  ( Brachydontes ) cowlitzensis  Weaver  and  Palmer, 
1922:16-17,  pi.  9,  fig.  19  [new  name  for  Modiola  ornata 
Gabb,  1864,  preoccupied], 

Brachidontes  cowlitzensis ? (Weaver  and  Palmer).  Stewart, 
1930:100-103,  pi.  8,  fig.  12. 

Brachidontes  cowlitzensis  I'M  caver  and  Palmer).  Turner,  1938: 
45-46,  pi.  6,  figs.  7-8.  Kleinpell  and  Weaver,  1963:197, 
pi.  29,  fig.  3.  Wolfe,  1977:3.  Givens  and  Kennedy,  1979: 
table  2. 

Volsella  (Brachidontes)  cowlitzensis  (Weaver  and  Palmer). 

Weaver,  1943:1 13-1 14,  pi.  26,  fig.  4. 

Brachidontes  ( Brachidontes ) cowlitzensis  (Weaver  and  Palm- 
er). Givens,  1974:43.  Squires,  1977,  table  1.  Moore,  1983: 
A66-A67,  pi.  17,  fig.  1. 

Primary  Type  Material.  ANSP  lectotype  4450  of  Modiola 
ornata  Gabb,  Domengine?  Formation,  Martinez,  California. 
CAS  holotype  7406  of  Modiolus  (Brachydontes)  ornatus 
Weaver  and  Palmer,  Cowlitz  Formation,  UW  locality  329. 

Molluscan  Stage  Range.  “Meganos”  through  lower  Oli- 
gocene  (Turritella  variata  lorenzana  Zone  of  Kleinpell  and 
Weaver,  1963). 

Geographic  Distribution.  San  Diego,  California  through 
Gulf  of  Alaska. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  374,  455,  466,  471d,  492,  514,  528. 
Shallow-marine  (regressive)  facies:  CSUN  locality  51 2d. 

Remarks.  Specimens  are  most  common  at  locality  371, 
where  they  are  fragile,  well-preserved  single  valves. 

Superfamily  Pinnacea 
Family  Pinnidae  Leach,  1819 
Genus  Pinna  Linne,  1758 

Type  Species.  By  subsequent  designation  (Children,  1823), 
Pinna  rudis  Linne,  1758. 

Pinna  lewisi  Waring,  1917 

Figure  lOf 

Pinna  lewisi  Waring,  1914:785;  1917:94,  pi.  15,  fig.  24. 
Pinna  (Pinna)  lewisi  Waring.  Moore,  1983:A79,  pi.  21,  fig. 
10. 

Primary  Type  Material.  CAS/SU  holotype  5194,  Llajas 
Formation,  SU  locality  2696. 


42  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  Simi  Valley  through  south  end 
of  San  Joaquin  Valley,  California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  548.  Shallow-marine  (regressive)  facies:  CSUN 
locality  475. 

Remarks.  Only  six  specimens  were  found,  and  five  are  from 
locality  475.  Preservation  is  mostly  as  internal  molds,  and 
four  specimens  are  articulated.  The  following  supplementary 
comments  are  based  on  an  examination  of  the  holotype.  It 
is  mostly  an  internal  mold  with  95  percent  of  the  shell  miss- 
ing. There  are  16  to  18,  very  closely  spaced  radial  ribs  and 
about  nine  comarginal  ribs  on  each  valve.  The  holotype  and 
the  Llajas  specimens  have  a median  sulcus. 

Pinna  llajasensis  Squires,  1983a 

Figure  lOg 

Pinna  n.  sp.  Vokes,  1939:50,  pi.  2,  fig.  14.  Moore,  1983:A79, 

pi.  22,  fig.  1. 

Pinna  llajasensis  Squires,  1983a:359-360,  fig.  21. 

Primary  Type  Material.  UCLA  holotype  59196,  Llajas 
Formation,  CSUN  locality  458. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  Simi  Valley  through  central  Cal- 
ifornia. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  374,  458. 

Remarks.  Only  two  specimens  were  found  and  these  are 
from  the  “Stewart  bed.”  Both  are  articulated,  incomplete 
specimens  (Squires,  1983).  The  holotype  is  the  figured  spec- 
imen (Fig.  lOg). 

P.  llajasensis  differs  from  P.  lewisi  in  having  fewer  radial 
ribs  (only  1 2)  which  are  more  widely  spaced.  P.  llajasensis 
also  has  no  comarginal  sculpture. 

In  the  original  description  by  Vokes  (1939)  of  his  Pinna 
n.  sp.,  he  mentioned  that  his  specimen  (UCMP  32595,  Do- 
mengine Formation,  UCMP  locality  2287)  had  23  ribs.  The 
following  supplementary  comments  are  based  on  an  exam- 
ination of  the  holotype.  There  are  12  ribs  per  single  valve 
and  about  23  for  the  entire  surface  including  both  valves. 

Order  Pterioida 
Suborder  Pteriina 
Superfamily  Pteriacea 
Family  Malleidae  Lamarck,  1819 
Genus  Nayadina  Munier-Chalmas,  1864 

Type  Species.  By  monotypy,  Nayadina  herberti  Munier- 
Chalmas,  1864. 

Subgenus  Exputens  Clark,  1934 

Type  Species.  By  subsequent  designation  (Vokes,  1939), 
Exputens  llajasensis  Clark,  1934. 


Nayadina  ( Exputens ) llajasensis  (Clark,  1934) 

Figures  lOh-i 

Exputens  llajasensis  Clark,  1934:270-271,  pi.  37,  figs.  1 1- 
18.  Vokes,  1939:51. 

Nayadina  ( Exputens ) llajasensis  (Clark).  Givens,  1974:44, 
pi.  1,  fig.  9.  Moore,  1983:A86-A87,  pi.  26,  figs.  10,  13. 

Primary  Type  Material.  UCMP  holotype  32391,  UCMP 
paratypes  32390,  32393,  Llajas  Formation,  UCMP  locality 
7004  = CSUN  locality  374. 

Molluscan  Stage  Range.  “Capay”  through  “Domengine.” 
Geographic  Distribution.  Simi  Valley  through  Pine  Moun- 
tain area,  southern  California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  374,  445,  447,  471b,  473,  479,  480. 

Remarks.  Specimens  occur  mainly  in  the  “Stewart  bed” 
at  and  in  the  vicinity  of  locality  374.  In  addition,  a few 
specimens  were  found  in  glauconitic  siltstone  at  locality  473, 
about  14  m above  the  “Stewart  bed”  at  the  type  section.  The 
bed  that  locality  473  occurs  in  probably  represents  a shallow- 
marine  (transgressive)  facies  bed  that  interfingers  with  the 
outer  shelf  and  slope  facies.  At  localities  374  and  473,  a few 
of  the  specimens  are  articulated. 

Superficially,  this  genus  may  resemble  Hiatella,  but  Elia- 
tella  is  a heterodont  with  two  weak  teeth  whereas  Nayadina 
lacks  hinge  teeth.  A view  of  the  hinge  of  N.  ( E .)  llajasensis 
is  given  in  Figure  lOi. 

Superfamily  Pectinacea 
Family  Spondylidae  Gray,  1826 
Genus  Spo ndylus  Linne,  1758 

Type  Species.  By  subsequent  designation  (Schmidt,  1818), 
Spondylus  gaederopus  Linne,  1758. 

Spondylus  carlosensis  Anderson,  1905 

Figure  lOj 

Spondylus  carlosensis  Anderson,  1905:194,  pi.  13,  fig.  1.  Ar- 
nold, 1 9 1 0:pl.  2,  figs.  6-7.  Dickerson,  191 5:pl.  1,  fig.  7. 
Anderson  and  Hanna,  1925: 189-190,  text  figure  10.  Vokes, 
1939:57,  pi.  3,  figs.  10,  13.  Kleinpell  and  Weaver,  1963: 
199,  pi.  31,  fig.  6. 

Primary  Type  Material.  CAS  holotype  56,  Domengine 
Formation,  west  and  north  of  Coalinga,  NW  Va  of  section  35, 
T 20  S,  R 14  E,  Fresno  County,  California. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  Simi  Valley  through  central  Cal- 
ifornia. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  374,  438,  444,  525. 

Remarks.  Specimens  are  relatively  uncommon  in  the  Lla- 
jas Formation,  and  they  consist  of  fragments  of  single  valves. 
The  most  complete  and  best  preserved  specimens  are  from 
locality  374. 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  43 


Figure  10.  Llajas  Formation  bivalves,  a.  Acila  ( Truncacila ) decisa  (Conrad,  1855),  right  valve,  LACMIP  hypotype  6554,  CSUN  loc.  467, 
x3.5.  b.  Nuculana  ( Saccella ) gabbi  (Gabb,  1864),  right  valve,  UCLA  hypotype  59279,  CSUN  loc.  467,  x3.  c.  Glycymeris  ( Glycymeris ) 
rosecanyonensis  Hanna,  1927,  right?  valve,  LACMIP  hypotype  6565,  CSUN  loc.  489,  x4.5.  d.  Glycymeris  (Glycymerita)  sagittata  (Gabb, 
1864),  right?  valve,  LACMIP  hypotype  6566,  CSUN  loc.  371,  x2.  e.  Brachidontes  ( Brachidontes ) cowlitzensis  (Weaver  and  Palmer,  1922), 
right  valve,  LACMIP  hypotype  6567,  CSUN  loc.  371,  x 1.5.  f.  Pinna  lewisi  (Waring,  1917),  internal  mold  of  right?  valve,  LACMIP  hypotype 


44  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Superfamily  Anomiacea 
Family  Anomiidae  Rafinesque,  1815 
Genus  Anomia  Linne,  1758 

Type  Species.  By  subsequent  designation  (Schmidt,  1818), 

Anomia  ephippium  Linne,  1758. 

Anomia  mcgoniglensis  Hanna,  1927 

Figure  10k 

Anomia  mcgoniglensis  Hanna,  1927:278,  pi.  31,  figs.  1,  2,  5, 
7.  Turner,  1938:46,  pi.  6,  figs.  4-6.  Weaver,  1943:100,  pi. 
22,  figs.  4-5.  Givens  and  Kennedy,  1976:974,  pi.  4,  figs. 
13-16.  Givens  and  Kennedy,  1979:table  2. 

Primary  Type  Material.  UCMP  syntypes  31009-31010, 
Delmar  Formation,  UCMP  locality  3981. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  San  Diego,  California  through 
southwestern  Oregon. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  438. 

Remarks.  Only  three  specimens  were  found  in  the  Llajas 
Formation.  They  occur  as  single  valves  unattached  to  any 
hard  substrate. 

Suborder  Ostreina 
Superfamily  Ostreacea 
Family  Ostreidae  Rafinesque,  1815 
Subfamily  Ostreinae  Rafinesque,  1815 
Genus  Ostrea  Linne,  1758 

Type  Species.  By  subsequent  designation  (ICZN  opin.  94 
and  356),  Ostrea  edulis  Linne,  1758. 

Ostrea  idriaensis  Gabb,  1869 

Figure  101 

Ostrea  idriaensis  Gabb,  1869:203,  pi.  33,  figs.  103b-d;  pi. 
34,  figs.  103,  103a.  Waring,  1917:78-79,  pi.  13,  fig.  10. 
Hanna,  1927:276,  pi.  30,  figs.  1-2;  pi.  31,  figs.  3-4.  Stewart, 
1930:126-127,  pi.  8,  fig.  3;  pi.  17,  fig.  1.  Vokes,  1935:291- 
304,  pi.  22-24.  Merriam  and  Turner,  1937:table  2.  Turner, 
1938:46,  pi.  6,  fig.  9.  Weaver,  1943:78-79,  pi.  15,  fig.  5. 
Givens,  1974:44.  Givens  and  Kennedy,  1 979:tables  2,  4. 
Not  Ostrea  haleyi  Hertlein,  1933:277-282,  pi.  18,  figs.  5-6. 


Primary  Type  Material.  MCZ  lectotype  15048,  Domen- 
gine Formation,  about  3 km  east  of  the  Hacienda  at  New 
Idria,  N V2  of  section  15,  T 17  S,  R 12  E,  Priest  Valley 
quadrangle,  San  Benito  County,  California. 

Molluscan  Stage  Range.  “Capay”  through  "Tejon.” 

Geographic  Distribution.  San  Diego,  California  through 
western  Washington. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  374,  444,  458,  468a,  478,  485,  548. 

Remarks.  Ostreid  fragments  occur  at  localities  502,  503, 
and  542  in  the  interfingering  coastal  alluvial-fan  facies  and 
shallow-marine  (transgressive)  facies,  but  generic  identifi- 
cations could  not  be  made.  Similarly,  ostreid  fragments  occur 
at  localities  463  and  526  in  the  lower  part  of  the  shallow- 
marine  (transgressive)  facies,  but  generic  identifications  could 
not  be  made.  Ostreid  fragments  and  single  valves  of  O.  id- 
riaensis are  fairly  common  in  the  uppermost  part  of  the 
“Stewart  bed.”  The  figured  specimen  (Fig.  101)  is  one  of  the 
largest  specimens. 

Subclass  Heterodonta 
Order  Veneroida 
Superfamily  Lucinacea 
Family  Lucinidae  Fleming,  1828 
Subfamily  Milthinae  Chavan,  1969 
Genus  Claibornites  Stewart,  1930 

Type  Species.  By  original  designation,  Lucina  rotunda  Lea, 
1833. 

Claibornites  diegoensis  (Dickerson,  1916) 

Figure  10m 

Lucina  diegoensis  Dickerson,  1916:484,  pi.  37,  figs.  la-b. 
Claibornites  diegoensis  ( Dickerson).  Givens,  1974:45-46,  pi. 

1,  fig.  15. 

Primary  Type  Material.  UCMP  holotype  1 1788,  Ardath 
Shale,  UCMP  locality  2226. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  San  Diego  through  Pine  Moun- 
tain area,  southern  California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  374,  445. 

Remarks.  At  locality  374,  only  two  single  valves  were  found. 
An  articulated  specimen  was  found  at  locality  445. 


6568,  CSUN  loc.  475,  x 1.  g.  Pinna  liajasensis  Squires,  1983a,  internal  mold  of  right  valve,  UCLA  holotype  59196,  CSUN  loc.  458,  x0.5. 
h and  i.  Navadina  (Exputens)  liajasensis  (Clark,  1934).  All  parts  from  CSUN  loc.  374.  h,  right  valve,  LACMIP  topotype  and  hypotype  6569, 
x 1.  i,  hinge  line,  LACMIP  topotype  and  hypotype  6570,  x 1.4.  j.  Spondylus  carlosensis  Anderson,  1905,  left  valve,  LACMIP  hypotype  6571, 
CSUN  loc.  374,  x l.  k.  Anomia  mcgoniglensis  Hanna,  1927,  right  valve,  UCLA  hypotype  59280,  CSUN  loc.  438,  x 1.5.  1.  Ostrea  idriaensis 
Gabb,  1869,  left  valve,  LACMIP  hypotype  6572,  CSUN  loc.  458,  x0.5.  m.  Claibornites  diegoensis  (Dickerson,  1916),  right  valve,  UCLA 
hypotype  59281,  CSUN  loc.  374,  x 1.  n and  o.  Venericardia  ( Pacifcor ) hornii  calafia  Stewart,  1930.  All  parts  from  CSUN  loc.  374  and  x0.5. 
n,  left  valve,  LACMIP  topotype  and  hypotype  6573.  o,  right  valve  hinge  line,  LACMIP  topotype  and  hypotype  6574.  p.  Venericardia  (Pacificor) 
aragonia  joaquinensis  (Vokes,  1939),  UCMP  hypotype  37433,  UCMP  loc.  7193  = CSUN  loc.  702,  x 1.  q and  r.  Glyptoactis  domenginica 
(Vokes,  1939),  LACMIP  hypotype  6575,  CSUN  loc.  371,  x2.5.  q,  left  valve  exterior,  r,  left  valve  interior. 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  45 


Superfamily  Carditacea 

Family  Carditidae  Fleming,  1 820 

Subfamily  Venericardiinae  Chavan,  1969 

Genus  Venericardia  Lamarck,  1801 

Type  Species.  By  subsequent  designation  (Schmidt,  1818), 
Venericardia  imbricata  Lamarck,  1801. 

Subgenus  Pacificor  Verastegui,  1953 

Type  Species.  By  original  designation,  Venericardia  ( Pa- 
cificor) mulleri  Verastegui,  1953. 

Venericardia  ( Pacificor ) hornii  calafia 
Stewart,  1930 

Figures  lOn-o 

Venericardia  hornii  calafia  Stewart,  1930:168-170,  pi.  11, 
fig.  2.  Turner,  1938:50,  pi.  14,  fig.  4.  Weaver,  1943:1 34— 
135,  pi.  28,  figs.  6-7;  pi.  31,  figs.  4-5. 

Venericardia  ( Pacificor ) calafia  Stewart.  Verastegui,  1953: 
28-30,  pi.  15,  figs.  3-5,  7;  pi.  16,  figs.  1-3;  pi.  17,  figs. 
1-2. 

Venericardia  ( Pacificor ) hornii  calafia  Stewart.  Givens,  1 974: 
47,  pi.  4,  fig.  1.  Saul,  1983:74,  76,  pi.  2,  figs.  9,  16-17. 

Primary  Type  Material.  UCMP  holotype  31450,  Llajas 
Formation,  UCMP  locality  7004  = CSUN  locality  374. 
Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  Simi  Valley,  California  through 
southwestern  Oregon. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  374,  445,  451,  458,  494,  548,  704. 

Remarks.  A growth  series  of  this  taxon  was  found  at  lo- 
cality 374.  At  this  locality,  most  of  the  specimens  are  single 
valves,  but  there  are  several  well-preserved,  large-sized  ar- 
ticulated specimens.  At  locality  548,  only  a single  articulated 
juvenile  was  found. 

Saul  (1983) considered  V.  (P.)  oregonensis  Verastequi  from 
the  Lookingglass  Formation  of  southwestern  Oregon  to  be 
an  immature  V.  (P.)  hornii  calafia.  V.  (P.)  hertleini  Veras- 
tequi is  also  probably  V.  (P.)  hornii  calafia  (Saul,  1984,  pers. 
commun.). 

Venericardia  ( Pacificor ) aragonia 
joaquinensis  {\ okes,  1939) 

Figure  lOp 

Venericardia  aragonia  var.  Turner,  1938:49,  pi.  13,  figs. 
6-9. 

Megacardita  ( Venericor ) hornii  joaquinensis  Vokes,  1939: 
69-70,  pi.  8,  figs.  1-2;  pi.  9,  figs.  1-2. 

Venericardia  ( Leuroactis ) schencki  Verastegui,  1953:50-51, 
pi.  4,  figs.  6-8. 

Venericardia  ( Leuroactis ) alisoensis  Verastegui,  1 953:52-53, 
pi.  10,  figs.  1-3. 

Venericardia  ( Leuroactis ) joaquinensis  (Vokes).  Verastegui, 
1953:60-61,  pi.  11,  figs.  1-4;  pi.  12,  figs.  4-6. 


Venericardia  (Leuroactis)  vokesi  Verastegui,  1953:61-62,  pi. 

14,  figs.  1-3. 

Venericardia  (Pacificor)  aragonia  joaquinensis  (V  okes).  Saul, 

1983:pl.  2,  figs.  7-8. 

Primary  Type  Material.  UCMP  holotype  15616,  Avenal 
Formation,  UCMP  locality  4170;  UCMP  paratype  15617, 
Avenal  Formation,  UCMP  locality  4169;  UCMP  paratype 
15618,  Avenal  Formation,  UCMP  locality  A-819. 

Molluscan  Stage  Range.  Uppermost  “Capay”  through 
“Domengine.” 

Geographic  Distribution.  Simi  Valley  through  central  Cal- 
ifornia. 

Local  Occurrence.  Interfingering  coastal  alluvial-fan  facies 
and  shallow-marine  (transgressive)  facies:  CSUN  localities 
452,  700,  702,  706. 

Remarks.  Most  specimens  in  the  Llajas  Formation  are 
fragments  of  single  valves.  Extraction  of  identifiable  speci- 
mens from  the  rock  is  usually  difficult.  At  locality  700,  only 
a single  juvenile  was  found. 

The  synonymy  of  this  taxon  is  based  on  work  by  Saul 
(1984,  pers.  commun.).  V.  (P.)  a.  smileyi  (Vokes)  may  be 
conspecific  with  V.  (P.  ) a.  joaquinensis,  but  more  taxonomic 
work  is  needed  to  resolve  the  matter  (Saul,  1 984,  pers.  com- 
mun.). 

The  figured  specimen  (Fig.  lOp)  is  UCMP  hypotype  37433 
from  UCMP  locality  7193  = CSUN  locality  702.  It  is  the 
same  specimen  as  used  by  Saul  (1983,  pi.  2,  fig.  8). 

Verastegui  (1953)  reported  V.  (L.)  schencki  from  the  Santa 
Susana  Formation,  but  his  vague  locality  information  makes 
it  impossible  to  know  exactly  where  he  collected  his  speci- 
mens. 

The  presence  of  V.  (P.)  a.  joaquinensis  in  the  zone  of  in- 
terfingering between  the  coastal  alluvial-fan  facies  and  the 
shallow-marine  (transgressive)  facies  of  the  Llajas  Formation 
refines  the  lower  limit  of  the  molluscan  stage  range  of  this 
taxon  as  uppermost  “Capay.” 

Subfamily  Carditesinae  Chavan,  1969 
Genus  Glyptoactis  Stewart,  1930 

Type  Species.  By  original  designation,  Venericardia  hadra 
Dali,  1903. 

Subgenus  Glyptoactis  s.s. 

Glyptoactis  ( Glyptoactis ) domenginica 
(Vokes,  1939) 

Figures  lOq-r 

Venericardia  ( Glyptoactis ?)  domenginica  Vokes,  1939:66,  pi. 

5,  figs.  7-9. 

Venericardia  (Glyptoactis)  domenginica  Vokes.  Verastegui, 

1953:43-44,  pi.  13,  fig.  1. 

Glyptoactis  domenginica  (Vokes).  Givens,  1974:47.  Squires, 

1977:table  1.  Givens  and  Kennedy,  1979:tables  1,  3. 

Primary  Type  Material.  UCMP  holotype  15611,  Domen- 
gine Formation,  UCMP  locality  A-1219;  UCMP  paratypes 
15612-15613,  Tejon  Formation,  UCMP  locality  A-1003. 


46  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Molluscan  Stage  Range.  “Domengine”  through  “Transi- 
tion.” 

Geographic  Distribution.  San  Diego  through  Mt.  Diablo, 
California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  507,  518,  529a. 

Remarks.  This  taxon  is  confined  to  the  shallow-marine 
(transgressive)  facies  of  the  Llajas  Formation.  Specimens  are 
most  abundant  at  locality  37 1 where  there  is  a growth  series. 
Preservation  is  excellent,  showing  the  angular  nodes  on  the 
ribs.  No  articulated  specimens  were  found. 

This  species  is  placed  in  the  subgenus  Glyptoactis  s.s.  be- 
cause the  radial  ribs  are  not  tripartite.  It  is  the  oldest  species 
of  this  subgenus. 

Superfamily  Crassatellacea 
Family  Crassatellidae  Ferussac,  1822 
Subfamily  Crassatellinae  Ferussac,  1822 
Genus  Crassatella  Lamarck,  1799 

Type  Species.  By  subsequent  designation  (Schmidt,  1818), 
Mactra  cygnaea  Lamarck,  1799  (not  Chemnitz,  1782)  [=C. 
gibba  Lamarck,  1801,  = Venus  ponderosa  Gmelin,  1791], 

Crassatella  uvasana  Conrad,  1855 

Figures  lla-g 

Crassatella  uvasana  Conrad,  1855:9;  1857,  pi.  2,  fig.  5.  Gabb, 
1864:214-215,  pi.  32,  fig.  284.  Stewart,  1930:141-143,  pi. 
12,  fig.  9.  Turner,  1938:47-48.  Givens,  1974:48. 
Crassatella  alta  Conrad,  1855:9;  1857:321  [not  Conrad,  1832: 
21,  pi.  7], 

Crassatella  grandis  Gabb,  1864:181,  pi.  24,  fig.  163;  1869: 
189. 

Astarte  semidentata  Cooper,  1894:48,  pi.  3,  figs.  44-45. 
Crassatellites  grandis  (Gabb).  Arnold,  1910:13,  pi.  2,  figs. 
10,  10a,  pi.  3,  fig.  14.  Dickerson,  1915:80,  pi.  1,  fig.  8;  pi. 
2,  figs,  la-b  [not  Waring,  1917:74,  pi.  12,  fig.  16  = Cras- 
satella branneri  fide  Nelson,  1925:410], 

Crassatellites  uvasana  (Conrad).  Arnold  and  Hannibal,  1913: 
569.  Dickerson,  1915:80,  pi.  2,  fig.  2.  Waring,  1917:59,  pi. 
8,  fig.  10. 

Crassatellites  mathewsonii  (Gabb).  Dickerson,  191 6:pl.  36, 
figs.  9a-b  (probably  C.  semidentata  (Cooper)  fide  Turner, 
1938:47-48). 

Crassatellites  uvasanus  (Conrad).  Anderson  and  Hanna,  1 925: 
172-174,  pi.  4,  figs.  2-3,  text  figure  7. 

Crassatellites  semidentata  (Cooper).  Hanna,  1927:282,  pi. 
35,  figs.  1-2. 

Crassatella  semidentata  (Cooper).  Turner,  1938:47-48. 
Crassatella  uvasana  semidentata  (Cooper).  Vokes,  1939:64- 
65,  pi.  4,  figs.  4,  6,  8,  10,  12.  Givens,  1974:48.  Squires, 
1977:table  1.  Givens  and  Kennedy,  1979:tables  1,  3. 
Crassatella  uvasana  uvasana  (Conrad).  Givens  and  Kenne- 
dy, 1979:table  4. 


Primary  Type  Material.  Holotype  undetected,  USNM  col- 
lection, Tejon  Formation,  Grapevine  Canyon,  Tejon  quad- 
rangle, Kern  County,  California. 

Molluscan  Stage  Range.  “Domengine”  through  “Tejon.” 

Geographic  Distribution.  San  Diego  through  central  Cal- 
ifornia. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  374,  444,  445,  447,  458,  471c,  481,  488, 
494.  Outer  shelf  and  slope  channel:  CSUN  locality  541. 

Remarks.  Ninety-six  specimens  were  found,  and  most  oc- 
cur as  single  valves.  Twelve  specimens  are  articulated.  A 
single,  abraded  fragment  was  found  at  the  base  of  a turbidite- 
filled  channel  at  locality  54 1 . Numerous  well-preserved  spec- 
imens of  C.  uvasana  occur  in  the  “Stewart  bed”  at  localities 
374  and  444.  Heights  range  from  20  to  90  mm  at  locality 
374  and  from  20  to  65  mm  at  locality  444.  There  is  a growth 
series  at  both  localities.  Many  of  the  juvenile  specimens  and 
a few  of  the  adult  specimens  are  articulated  at  both  localities. 
For  the  disarticulated  specimens,  the  ratio  of  right  valves  to 
left  valves  is  approximately  1:1.  Based  on  criteria  listed  by 
Fagerstrom  ( 1 964),  the  C.  uvasana  specimens  at  these  “Stew- 
art bed”  localities  are  clearly  in  situ. 

There  is  a gradual  change  in  the  external  appearance  of 
the  shells  from  juvenile  to  adult  individuals  at  these  localities 
(Figs.  1 la-c).  The  juvenile  specimens  have  higher  and  more 
prominent  beaks,  a less  deep  lunule,  a less  steep  posterior 
dorsal  slope,  a more  concave  anterior  dorsal  margin,  and  a 
less  obvious  escutcheon  on  the  left  valve.  The  adult  speci- 
mens have  a more  triangular  shape  and  lower,  more  rounded 
beaks.  The  escutcheon  on  the  right  valve  is  much  larger  than 
that  on  the  left.  Intermediate-sized  specimens  (with  a height 
of  about  45  mm)  are  transitional  between  the  two. 

The  comarginal  ribbing  is  identical  in  both  the  juvenile 
and  adult  specimens.  There  are  numerous  very  fine,  closely 
spaced  comarginal  ribs.  At  regular  intervals  there  are  comar- 
ginal rugae.  Upon  decortication,  especially  in  the  beak  areas, 
shell  between  the  rugae  has  a tendency  to  chip  off,  leaving 
depressed  areas  outlined  by  the  rugae.  Only  the  adult  spec- 
imens show  cancellate  sculpture  along  the  ventral  margins 
of  the  valves. 

Interior  shell  features  of  juvenile  and  adult  specimens  of 
C.  uvasana  are  identical  (Figs.  1 1 d— g).  The  resilifer  extends 
halfway  to  the  lower  margin  of  the  hinge  plate  and  the  pit  is 
large.  The  anterior  muscle  scar  is  reniform,  and  the  pallial 
line  is  deeply  impressed.  In  both  juvenile  and  adult  speci- 
mens the  right  valve  has  the  following  features:  anterior  ven- 
tral margin  of  the  hinge  plate  tends  to  be  swollen  and  fits 
into  a socket  in  the  left  valve;  anterior  cardinal  is  large, 
wedge-shaped,  vertically  below  the  beak,  its  upper  end  mar- 
gining part  of  the  resilifer;  the  floor  of  the  lunule  tends  to 
wrap  around  the  anterior  cardinal  in  some  specimens  (Fig. 

1 If);  posterior  cardinal  is  weak;  edge  of  the  escutcheon  is 
swollen  and  projects  as  a long  ridge  that  fits  into  a socket  in 
the  left  valve. 

In  both  juvenile  and  adult  specimens  of  C.  uvasana.  the 
left  valve  has  the  following  features:  posterior  ventral  margin 
of  the  hinge  plate  is  slightly  swollen;  anterior  and  posterior 
cardinals  are  about  equal  in  size;  floor  of  lunule  tends  to  wrap 


Contributions  in  Science,  Number  350 


Squires:  Simni  Valley  Eocene  Mollusks  47 


Figure  11.  Llajas  Formation  bivalves  (continued),  a through  g.  Crassatella  uvasana  Conrad,  1855.  All  parts  from  CSUN  loc.  374.  a,  left 
valve,  LACMIP  hypotype  6576,  x 1.  b,  left  valve,  LACMIP  hypotype  6577,  x 1.  c,  left  valve,  LACMIP  hypotype  6578,  x 1.  d,  right  valve 
interior  of  juvenile,  LACMIP  hypotype  6579,  x 1.  e,  left  valve  hinge  line  of  juvenile,  LACMIP  hypotype  6580,  x 1.5.  f,  right  valve  hinge  line 
of  adult,  LACMIP  hypotype  6581,  xl.  g,  left  valve  interior  of  adult,  LACMIP  hypotype  6582,  x 1. 


48  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


around  the  anterior  cardinal  in  some  specimens,  ventral  mar- 
gin of  lunule  tends  to  be  swollen.  In  the  left  valve,  as  well  as 
in  the  right  valve,  the  posterior  part  of  the  hinge  plate  is 
largely  occupied  by  a smooth  surface  without  hinge  struc- 
tures. 

If  it  had  not  been  for  the  presence  of  a growth  series,  the 
juvenile  specimens  of  C.  uvasana  would  have  been  identified 
as  C.  uvasana  semidentata  (Cooper),  and  the  adult  specimens 
would  have  been  identified  as  C.  uvasana.  Based  on  the 
transition  of  shell  characters  from  the  juvenile  and  adult,  as 
well  as  the  identical  shell  characters  mentioned  above,  it  is 
concluded  that  C.  uvasana  semidentata  is  an  unjustified  sub- 
species. 

According  to  Hanna  ( 1 927)  and  Givens  ( 1 974),  the  nature 
of  the  beaks  is  the  main  distinguishing  character  between  C. 
uvasana  and  C.  uvasana  semidentata.  In  actuality,  this  char- 
acter can  be  used  to  distinguish  juvenile  from  adult  speci- 
mens of  C.  uvasana  as  the  beaks  become  more  incurved  with 
age.  It  is  interesting  to  note  that  if  a juvenile  specimen  is 
placed  on  a flat  surface,  viewed  laterally,  and  elevated  along 
the  venter  (simulating  additional  shell  material),  the  beak 
area  then  looks  identical  to  that  of  an  adult  specimen. 

The  molluscan  stage  range  of  C.  u.  semidentata  had  been 
reported  as  “Domengine”  through  “Transition”  and  that  of 
C.  uvasana  as  “Tejon”  (Givens,  1 974).  Because  the  two  taxa 
are  identical,  the  molluscan  stage  range  of  C.  uvasana  can 
be  extended  to  “Domengine”  through  “Tejon.” 

Superfamily  Cardiacea 

Family  Cardiidae  Lamarck,  1 809 

Subfamily  Cardiinae  Lamarck,  1809 

Genus  Acanthocardia  Gray,  1851 

Type  Species.  By  subsequent  designation  (Stoliczka,  1 870), 
Cardium  aculeatum  Linne,  1758. 

Subgenus  Schedocardia  Stewart,  1930 

Type  Species.  By  original  designation,  Cardium  hatche- 
tigbeense  Aldrich,  1886. 

Acanthocardia  ( Schedocardia ) brewerii 
(Gabb,  1864) 

Figures  12a-b 

Cardium  brewerii  Gabb,  1864:173,  pi.  24,  fig.  155.  Arnold, 

1 907 :pl.  39,  fig.  5.  McLaughlin  and  Waring,  191 5:fig.  14. 
Waring,  1 9 1 7:pl.  14,  fig.  9.  Anderson  and  Hanna,  1925: 
165-166,  pi.  1,  fig.  3.  Clark,  1929:pl.  12,  fig.  7. 
Plagiocardium  ( Schedocardia ) brewerii  (Gabb).  Stewart,  1 930: 
256-258,  pi.  12,  fig.  6.  Turner,  1938:52-53,  pi.  9,  figs.  6- 
7.  Vokes,  1939:75,  pi.  1 1,  figs.  1-4.  Stewart,  1946:pl.  I 1, 
fig.  20. 

Plagiocardium  brewerii  (Gabb).  Merriam  and  Turner,  1937: 
table  2. 

Loxocardium (Schedocardia)  brewerii) Gabb).  Weaver,  1943: 
153-154,  pi.  35,  figs.  15,  16,  18;  pi.  38,  figs.  1,  9;  pi.  104, 
fig.  12. 


Cardium  ( Trachycardium)  brewerii  brewerii  (Gabb).  Klein- 
pell  and  Weaver,  1963:201-202,  pi.  34,  figs.  1-2. 
Acanthocardia  (Schedocardia)  brewerii  (Gabb).  Givens,  1974: 
48-49,  pi.  1,  fig.  17. 

Acanthocardia  brewerii  (Gabb).  Givens  and  Kennedy,  1979: 
table  4. 

Primary  Type  Material.  ANSP  lectotype  4560,  Tejon  For- 
mation, east  of  north  end  of  Grapevine  Canyon,  Kern  Coun- 
ty, California. 

Molluscan  Stage  Range.  “Capay”  through  “Tejon.” 
Geographic  Distribution.  San  Diego,  California  through 
southwestern  Washington. 

Local  Occurrence.  Interfingering  coastal  alluvial-fan  facies 
and  shallow-marine  (transgressive)  facies:  CSUN  localities 
545,  702.  Shallow-marine  (transgressive)  facies:  CSUN  lo- 
calities 371,  374,  548,  704. 

Remarks.  This  taxon  is  somewhat  uncommon  in  the  Llajas 
Formation.  It  is  most  common  at  locality  374  where  it  occurs 
as  single  valves.  The  largest  specimens  are  also  from  this 
locality. 

Subfamily  Protocardiinae  Keen,  1951 

Genus  Nemocardium  Meek,  1876 

Type  Species.  By  subsequent  designation  (Sacco,  1899), 
Cardium  semiasperum  Deshayes,  1858. 

Nemocardium  linteum  (Conrad,  1855) 

Figure  1 2c 

Cardium  linteum  Conrad,  1855:3,  9;  1857 :pl.  2,  fig.  1.  An- 
derson and  Hanna,  1925:166-167,  pi.  3,  fig.  3. 

Cardium  cooperii  Gabb,  1864:172,  pi.  24,  figs.  154,  154a. 
Arnold,  1 907 :pl  38,  figs.  2-2a.  Waring,  191 7:pl.  13,  fig. 
3.  Hanna,  1927:285,  pi.  41,  figs.  6-7. 

Cardium  dalh  Dickerson,  1913:289,  pi.  14,  figs.  4a-c. 
Cardium  marysvillensis  Dickerson,  1916:482  [new  name  for 
Cardium  dalli  Dickerson,  1913,  preoccupied]. 

Cardium  (Protocardium)  marysvillensis  Dickerson.  Clark  and 
Woodford,  1927:94,  pi.  15,  fig.  12. 

Nemocardium  linteum  (Conrad).  Stewart,  1930:275-277,  pi. 
8,  fig.  6.  Turner,  1938:52,  pi.  10,  fig.  10.  Vokes,  1939:7 6— 
77,  pi.  11,  figs.  6,  9.  Weaver,  1943:159-160,  pi.  38,  fig.  3; 
1953:28.  Stewart,  1946:pl.  11,  fig.  19.  Zinsmeister,  1974: 
97-98,  pi.  9,  figs.  7-9;  1 983a:pl.  2,  fig.  7.  Givens  and  Ken- 
nedy, 1 979:table  4. 

Cardium  (Nemocardium)  linteum  Conrad.  Kleinpell  and 
Weaver,  1963:202,  pi.  34,  fig.  4. 

Nemocardium  (Nemocardium)  linteum  (Conrad).  Givens, 
1974:49.  Squires,  1977:table  1. 

Primary  Type  Material.  USNM  holotype  1834,  Domen- 
gine Formation,  near  Martinez,  California. 

Molluscan  Stage  Range.  “Martinez”  through  “Tejon.” 
Geographic  Distribution.  San  Diego,  California  through 
southwestern  Oregon. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  373,  374,  438,  444,  445,  458,  479,  488,  493, 
513,  525,  548. 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  49 


Remarks.  This  taxon  is  most  common  in  the  upper  part 
of  the  shallow-marine  (transgressive)  facies.  At  locality  438, 
six  specimens  were  found,  one  of  which  was  articulated.  In 
the  “Stewart  bed”  at  locality  374,  21  specimens  were  found 
and  one  was  articulated. 

Superfamily  Solenacea 
Family  Solenidae  Lamarck,  1809 
Genus  Solena  Morch,  1853 

T ype  Species.  By  subsequent  designation  (Stoliczka,  1871), 
Solen  obliquus  Spengler,  1794. 

Subgenus  Eosolen  Stewart,  1930 

Type  Species.  By  original  designation,  Solen  plagiaulax 
Cossmann,  1906. 

Solena  ( Eosolen ) novacularis 
(Anderson  and  Hanna,  1928) 

Figure  12d 

Solen  novacula  Anderson  and  Hanna,  1925:147,  pi.  6,  fig.  9. 

Hanna,  1927:294,  pi.  43,  fig.  1. 

Not  Solen  novacula  Montagu,  1803:47. 

Solen  novacularis  Anderson  and  Hanna,  1928:65-66  [new 
name  for  Solen  novacula  Anderson  and  Hanna,  1925, 
preoccupied], 

Solena  ( Eosolen ) coosensis  Turner,  1938:62-63,  pi.  9,  figs. 
1-2.  Vokes,  1939:96,  pi.  15,  fig.  5.  Givens,  1974:49-50, 
pi-  2,  fig.  1. 

Solena  coosensis  Turner.  Weaver,  1943:229,  pi.  52,  fig.  16; 
pi.  53,  fig.  13. 

Solena  novacularis  (Anderson  and  Hanna).  Givens  and  Ken- 
nedy, 1979:table  4. 

Primary  Type  Material.  CAS  holotype  882  of  Solen  no- 
vacula Anderson  and  Hanna  and  of  Solen  novacularis  An- 
derson and  Hanna,  Tejon  Formation,  CAS  locality  792. 

Molluscan  Stage  Range.  Uppermost  “Capay”?,  “Domen- 
gine”  through  “Tejon.” 

Geographic  Distribution.  San  Diego,  California  through 
southwestern  Oregon. 

Local  Occurrence.  Interfingering  coastal  alluvial-fan  facies 
and  shallow-marine  (transgressive)  facies:  CSUN  locality  701? 
Shallow-marine  (transgressive)  facies:  CSUN  localities  374, 
444,  445,  449,  458,  477,  494,  500. 

Remarks.  This  species  helps  to  characterize  the  “Stewart 
bed.”  At  many  localities  of  the  “Stewart  bed,”  and  especially 
at  locality  374,  specimens  are  fairly  common  and  many  are 
articulated.  A solenid  at  locality  701  may  be  this  species. 

A previously  assigned  “Capay”  age  (Turner,  1 938;  Givens, 
1974)  for  the  lower  range  limit  of  this  species  was  based  on 
occurrence  in  the  “upper  Umpqua”  Formation  of  south- 
western Oregon.  The  “upper  Umpqua,”  however,  is  actually 
equivalent  to  both  the  “Capay”  and  “Domengine”  “Stages” 
(Baldwin,  1974;  Miles,  1981).  A lower  range  limit  of  upper- 
most “Capay,”  nevertheless,  is  apparent  because  of  the  pos- 
sible occurrence  of  this  species  in  the  zone  of  interfingering 


between  the  coastal  alluvial-fan  facies  and  the  shallow-ma- 
rine (transgressive)  facies  of  the  Llajas  Formation. 

Superfamily  Tellinacea 
Family  Tellinidae  Blainville,  1814 
Subfamily  Macominae  Olsson,  1961 
Genus  Macoma  Leach,  1819 

Type  Species.  By  monotypy,  Macoma  tenera  Leach,  1819 
[=Tellina  calcarea  Gmelin,  1791]. 

Macoma  rosa  Hanna,  1927 

Figure  12e 

Macoma  rosa  Hanna,  1927:292,  pi.  41,  figs.  2-5,  8.  Clark, 

1929:pl.  6,  fig.  15.  Givens  and  Kennedy,  1979:table  1. 

Primary  Type  Material.  UCMP  holotype  3101)4,  Ardath 
Shale,  UCMP  locality  3993;  UCMP  paratype  31095,  Ardath 
Shale,  UCMP  locality  5089;  UCMP  paratypes  3 1 096-3 1 097, 
Ardath  Shale,  UCMP  locality  5085. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  San  Diego  through  Simi  Valley, 
California. 

Local  Occurrence.  Shallow-marine  (regressive)  facies: 
CSUN  localities  462,  489,  512a,  51 2d,  544. 

Remarks.  This  taxon  helps  to  characterize  the  shallow- 
marine  (regressive)  facies.  Specimens  are  usually  few  in  num- 
ber and  preserved  as  external  and  internal  molds  of  articu- 
lated individuals. 

Family  Psammobiidae  Fleming,  1828 
Subfamily  Psammobiinae  Fleming,  1828 
Genus  Gari  Schumacher,  1817 

Type  Species.  Pending  decision  by  the  ICZN,  Gari  vulgaris 
Schumacher,  1817  ( =Solen  amethystus  Wood,  1818). 

Gari  cf.  G.  eoundulata  Vokes,  1939 

Figure  12f 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  374,  47 Id. 

Remarks.  Specimens  are  relatively  rare  in  the  Llajas  For- 
mation. They  are  preserved  as  internal  molds  of  single  valves, 
making  species  identification  impossible. 

Superfamily  Veneracea 
Family  Veneridae  Rafinesque,  1815 
Subfamily  Pitarinae  Stewart,  1930 
Genus  Callista  Poli,  1791 

Type  Species.  By  subsequent  designation  (Meek,  1876), 
Venus  chione  Linne,  1758. 


50  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Subgenus  Costacallista  Palmer,  1927 

Type  Species.  By  original  designation,  Venus  erycina  Linne, 
1758. 

Callista  ( Costacallista ) cf. 

C.  (C.)  hornii  (Gabb,  1864) 

Figure  12g 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  373,  465,  466,  467,  704. 

Remarks.  A few  specimens  were  found  as  single  valves. 
Hinge-line  details  were  imperfectly  preserved,  hence  a certain 
species  identification  could  not  be  made. 

Genus  Callocardia  A.  Adams,  1864 

Type  Species.  By  monotypy,  Callocardia  guttata  A.  Adams, 
1864. 

Subgenus  Nitidavenus  Vokes,  1939 

Type  Species.  By  original  designation,  Cytherea  nitida 
Deshayes,  1858. 

Callocardia  ( Nitidavenus ) tejonensis 
(Waring,  1914) 

Figure  12h 

Isocardia  tejonensis  Waring,  1914:784-785;  1917:93,  pi.  15, 
fig.  14. 

cf.  “ Isocardia  tejonensis"  Waring.  Turner,  1938:58,  pi.  11, 
figs.  1-4. 

Nitidavenus  tejonensis  (Waring).  Vokes,  1 939:83-84,  pi.  12, 
figs.  11,  13-16. 

Primary  Type  Material.  SU  holotype  189,  SU  paratypes 
5188-5190,  Llajas  Formation,  SU  locality  2696. 

Molluscan  Stage  Range.  “Capay”  through  “Domengine.” 
Geographic  Distribution.  Simi  Valley,  California  through 
southwestern  Oregon. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  372. 

Remarks.  Only  four  single  valves  were  found  in  the  Llajas 
Formation. 

Genus  Pi  tar  Romer,  1857 
Type  Species.  By  monotypy,  Venus  lumens  Gmelin,  1791. 
Subgenus  Calpitaria  Jukes-Browne,  1908 

Type  Species.  By  original  designation,  Cytherea  sulcataria 
Deshayes,  1824. 

Pitar  (Calpitaria)  uvasanus  (Conrad,  1855). 

Figures  12i-j 

Meretrix  uvasana  Conrad,  1855:9;  1857:pl.  2,  fig.  3.  Gabb, 
1864:163-164,  pi.  30,  fig.  248. 

Meretrix  tejonensis  Dickerson,  1915:53-54,  pi.  3,  figs.  3a-b, 
not  2a-b.  [Unjustified  new  name  for  Meretrix  uvasana 
Conrad  of  Gabb,  1864,  fide  Anderson  and  Hanna,  1925: 
160-161  and  Stewart,  1930:236.] 


Not  Pitaria  tejonensis  (Dickerson).  Anderson  and  Hanna, 
1925:160-161,  pi.  3,  fig.  5. 

Pitaria  uvasana  (Conrad).  Anderson  and  Hanna,  1925:161, 
pi.  5,  figs.  3-4. 

Pitar  (Calpitaria)  uvasanus  (Conrad).  Stewart,  1930:235-236, 
pi.  12,  fig.  7.  Weaver,  1943:178-179,  pi.  47,  fig.  13.  Givens, 
1974:53. 

Pitar  uvasanus  uvasanus  (Conrad).  Givens  and  Kennedy, 
1979:tables  3-4. 

Primary  Type  Material.  ANSP  neotype  4457,  Tejon  For- 
mation, Grapevine  Canyon,  Kern  County,  California. 
Molluscan  Stage  Range.  “Domengine”  through  “Tejon.” 
Geographic  Distribution.  San  Diego,  California  through 
western  Oregon. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  371,  374. 

Remarks.  Specimens  are  fairly  uncommon  and  generally 
consist  of  fragments  of  single  valves.  The  occurrence  of  this 
taxon  in  the  Llajas  Formation  extends  its  range  into  the 
“Domengine  Stage.”  Previously,  it  had  been  reported  as 
ranging  from  the  “Transition”  through  the  “Tejon”  (Givens, 
1974). 

Meretrix  tejonensis  Dickerson,  1915:53,  pi.  3,  figs.  2a-b  = 
Pitar  ( Lamelliconcha ) dickersoni  Givens,  fide  Givens,  1974: 
53-54. 


Subgenus  Lamelliconcha  Dali,  1902 

Type  Species.  By  original  designation,  Cytherea  concinna 
J.  Sowerby,  1835a. 

Pitar  ( Lamelliconcha ) joaquinensis 
Vokes,  1939 

Figure  12k 

Meretrix  hornii  Gabb.  Arnold,  1 9 10:pl.  3,  fig.  9.  [Misiden- 
tification.] 

Pitar  (Lamelliconcha)  joaquinensis  Vokes,  1939:85-86,  pi. 

13,  figs.  9-12.  Givens,  1974:54,  pi.  3,  fig.  7. 

Pitar ? joaquinensis  Vokes.  Stewart,  1946:pl.  12,  fig.  12. 
Pitar  joaquinensis  Vokes.  Givens  and  Kennedy,  1979: 
table  1. 

Primary  Type  Material.  UCMP  holotype  15674,  Domen- 
gine Formation,  UCMP  locality  A- 1027;  UCMP  paratype 
1 5675,  Domengine?  Formation,  UCMP  locality  4175;  UCMP 
paratype  15676,  Domengine  Formation,  UCMP  locality 
A- 1027;  UCMP  paratype  1 5677,  Avenal  Formation,  UCMP 
locality  A- 1280. 

Molluscan  Stage  Range.  “Domengine.” 

Geographic  Distribution.  Simi  Valley  through  central  Cal- 
ifornia. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  371. 

Remarks.  Only  a few  single  valves  were  found  in  the  Llajas 
Formation. 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  51 


Figure  12.  Llajas  Formation  bivalves  (continued),  a and  b.  Acanthocardia  (Schedocardia ) breweri  (Gabb,  1864),  right  valve,  LACMIP  hypotype 

6583,  CSUN  loc.  371,  x2.  a,  exterior,  b,  interior,  c.  Nemocardium  linteum  (Conrad,  1855),  internal  mold  of  right?  valve,  LACMIP  hypotype 

6584,  CSUN  loc.  374,  x 1.  d.  Solena  ( Eosolen ) novacularis  (Anderson  and  Hanna,  1928),  left  valve,  LACMIP  hypotype  6585,  CSUN  loc. 
374,  x l.  e.  Macoma  rosa  Hanna,  1927,  internal  mold  of  right  valve,  LACMIP  hypotype  6586,  CSUN  loc.  489,  x 1.5.  f.  Gari  cf.  G.  eoundulata 
Vokes,  1939,  internal  mold  of  right  valve,  LACMIP  hypotype  6587,  CSUN  loc.  47  Id,  x 1.  g.  Callista  ( Costacallista ) cf.  C.  (C.)  hornii  (Gabb, 
1864),  right  valve,  LACMIP  hypotype  6588,  CSUN  loc.  465,  x2.  h.  Callocardia  (Nitidavenus)  tejonensis  (Waring,  1914),  left  valve,  LACMIP 


52  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Subfamily  Tapetinae  H.  and  A.  Adams,  1857 

Genus  Marcia  H.  and  A.  Adams,  1857 

Type  Species.  By  subsequent  designation  (Kobelt,  1883), 
“ Venus  pinguis  Chemnitz”  [=  V.  opima  Gmelin,  1791], 

Subgenus  Mercimonia  Dali,  1902 

Type  Species.  By  original  designation,  Venus  bernayi  Coss- 
mann,  1888. 

Marcia  ( Mercimonia ) bunkeri  (Hanna,  1927) 

Figure  121 

Dosinia  bunkeri  Hanna,  1927:287,  pi.  42,  figs.  4,  6. 
Mercimonia  bunkeri  (Hanna).  Turner,  1938:60,  pi.  10,  figs. 
5-9.  Vokes,  1939:77,  pi.  11,  fig.  12.  Givens  and  Kennedy, 
1979:87,  table  3. 

Marcia  ( Mercimonia ) bunkeri  (Hanna).  Weaver,  1943:192, 
pi.  45,  fig.  5;  pi.  47,  figs.  8,  11. 

Primary  Type  Material.  UCMP  holotype  30950,  Ardath 
Shale,  UCMP  locality  5069;  UCMP  paratype  30951,  Ardath 
Shale,  UCMP  locality  3976. 

Molluscan  Stage  Range.  “Domengine”  through  “Transi- 
tion.” 

Geographic  Distribution.  San  Diego,  California  through 
southwestern  Oregon. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  374. 

Remarks.  Only  three  specimens  were  found  in  the  Llajas 
Formation,  and  of  these,  two  are  articulated. 

Order  Myoida 
Suborder  Myina 
Superfamily  Myacea 
Family  Corbulidae  Lamarck,  1818 
Subfamily  Corbulinae  Lamarck,  1818 
Genus  Corbula  Bruguiere,  1797 

Type  Species.  By  subsequent  designation  (Schmidt,  1818), 
Corbula  sulcata  Lamarck,  1801. 

Subgenus  C laryocorbula  Gardner,  1926 

Type  Species.  By  original  designation,  Corbula  alaba- 
miensis  Lea,  1833. 


Corbula  ( Caryocorbula ) dickersoni 
(Weaver  and  Palmer,  1922) 

Figure  12m 

Corbula  dickersoni  Weaver  and  Palmer,  1922:24-25,  pi.  9, 
figs.  9-10.  Clark,  1938:700,  pi.  1,  fig.  17.  Weaver,  1943: 
257-258,  pi.  61,  figs.  13,  16-17,  20. 

Corbula  ( Caryocorbula ) dickersoni  Weaver  and  Palmer. 
Vokes,  1939:98,  pi.  16,  figs.  1,  5,  9.  Givens,  1974:57,  pi. 
4,  fig.  7. 

Primary  Type  Material.  CAS  holotype  7452,  CAS  syn- 
types  7452A-B,  Cowlitz  Formation,  UW  locality  329. 

Molluscan  Stage  Range.  Uppermost  “Capay”  through 
“Tejon.” 

Geographic  Distribution.  Simi  Valley,  California  through 
southwestern  Washington. 

Local  Occurrence.  Interfingering  coastal  alluvial-fan  facies 
and  shallow-marine  (transgressive)  facies:  CSUN  locality  545. 
Shallow-marine  (transgressive)  facies:  CSUN  localities  371, 

373,  455,  484,  498,  501,  507,  540.  Shallow-marine  (regres- 
sive) facies:  CSUN  locality  475. 

Remarks.  This  taxon  is  most  common  at  locality  371, 
where  the  specimens  are  similar-sized  single  valves. 

The  presence  of  C.  (C.)  dickersoni  in  the  zone  of  interfin- 
gering between  the  coastal  alluvial-fan  facies  and  the  shallow- 
marine  (transgressive)  facies  of  the  Llajas  Formation  extends 
the  molluscan  stage  range  of  this  taxon  into  the  uppermost 
“Capay.”  Previously,  the  lower  range  limit  had  been  reported 
as  the  “Domengine”  (Vokes,  1939;  Givens,  1974). 

Suborder  Phoiadina 

Superfamily  Pholadacea 

Family  Teredinidae  Rafinesque,  1815 

Subfamily  Teredininae  Rafinesque,  1815 

Genus  Teredo  Linne,  1758 

Type  Species.  By  subsequent  designation  (ICZN,  1926, 
opin.  94),  Teredo  navalis  Linne,  1758. 

Teredol  sp. 

Figure  12n 

Local  Occurrence.  Interfingering  coastal  alluvial-fan  facies 
and  shallow-marine  (transgressive)  facies:  CSUN  locality 
469c.  Shallow-marine  (transgressive)  facies:  CSUN  localities 

374,  438,  444,  469,  470a,  488,  534.  Outer  shelf  and  slope 


hypotype  6589,  CSUN  loc.  371,  x 1.  i and  j.  Pitar  ( Calpitaria)  uvasanus  (Conrad,  1855),  left  valve,  LACMIP  hypotype  6590,  CSUN  loc.  371, 
x 1.5.  i,  exterior,  j,  interior,  k.  Pitar  ( La melliconcha)  joaquinensis  Vokes,  1939,  partial  specimen,  right  valve,  LACMIP  hypotype  6591,  CSUN 
loc.  371,  x l . l.  Marcia  ( Mercimonia ) bunkeri  ( Hanna,  1927),  left?  valve,  UCLA  hypotype  59282,  CSUN  loc.  374,  x 1 . m.  Corbula  (Caryocorbula) 
dickersoni  (Weaver  and  Palmer,  1922),  right  valve,  LACMIP  hypotype  6592,  CSUN  loc.  371,  x 2.75.  n.  Teredo ? sp.,  in  petrified  wood, 
LACMIP  hypotype  6593,  CSUN  locality  524,  x0.5.  o.  Cardiomya  aff.  C.  russelli  (Hanna,  1927),  internal  mold  of  right  valve,  LACMIP 
hypotype  6594,  CSUN  loc.  51 2d,  x4. 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  53 


Figure  13.  Llajas  Formation  nautiloid,  crab,  spatangoid,  and  shark  teeth,  a.  Aturia  myrlae  Hanna,  1927,  lateral  view,  LACMIP  hypotype 
6595,  CSUN  loc.  374,  xl.75.  b.  Unidentifiable  spirulimorph  sepiid,  ventral  view  of  partial  phragmocone,  UCLA  hypotype  59197,  CSUN 
loc.  493,  x 2.6.  c.  Glyphithyreus  weaveri  (Rathbun,  1926),  carapace  and  portion  of  right  cheliped,  dorsal  view,  UCLA  hypotype  59273,  CSUN 
loc.  548,  x l . d.  Schizaster  diabloensis  Kew,  1920,  internal  mold,  dorsal  view,  LACMIP  hypotype  6596,  CSUN  loc.  482,  x 1.  e.  Odontaspis 
sp.,  LACMIP  hypotype  6597,  CSUN  loc.  374,  x 1.  f.  Isurus  cf.  I.  praecursor  (Leriche,  1906),  LACMIP  hypotype  6598,  CSUN  loc.  371,  x3. 


facies:  CSUN  locality  446.  Outer  shelf  and  slope  channel 
facies:  CSUN  locality  541. 

Remarks.  Teredo ? sp.  occurs  in  pieces  of  petrified  wood. 
In  most  cases.  Teredo ? sp.  is  represented  by  calcareous-lined 
burrows,  but  unlined  burrows  can  be  present  in  the  same 
piece  of  wood.  Generic  determination  is  uncertain,  and  it  is 
very  possible  that  future  workers  will  assign  these  fossils  to 
another  genus.  Tertiary  teredinids  are  in  need  of  much  taxo- 
nomic work. 

Subclass  Anomalodesmata 
Order  Pholadomyoida 
Superfamily  Poromyacea 
Family  Cuspidariidae  Dali,  1886 
Genus  Cardiomya  A.  Adams,  1864 

Type  Species.  By  monotypy,  Neaera  gouldiana  Hinds, 
1843. 

Cardiomya  aff.  C.  russelli  (Hanna,  1927) 

Figure  12o 

Local  Occurrence.  Shallow-marine  (regressive)  facies: 
CSUN  locality  5 1 2d. 


Remarks.  Only  a single  specimen  was  found,  and  it  is  an 
internal  mold  of  a right  valve.  The  specimen  agrees  closely 
with  the  description  of  C.  russelli  given  by  Hanna  (1927: 
280-28 1 ) in  possessing  radial  ribs  and  a long,  straight  smooth 
rostrum.  The  Llajas  specimen  differs  in  that  it  has  at  least 
19  radial  ribs  rather  than  16. 

Class  Cephalopoda 
Subclass  Nautiloidea 
Order  Nautilida 
Superfamily  Nautilacea 
Family  Aturiidae  Hyatt,  1894 
Genus  Aturia  Bronn,  1838 

Type  Species.  By  subsequent  designation  (Herrmannsen, 
1846),  Nautilus  aturi  Basterot,  1825,  and  virtual  tautonymy. 

Aturia  myrlae  Hanna,  1927 

Figure  13a 

Aturia  myrli  Hanna,  1927:331,  pi.  57,  figs.  1,  6. 

Aturia  myrlae  Hanna.  Schenck,  1931:454-456,  pis.  67-68; 
text  figures  4-3,  4-19.  Yokes,  1939:107,  pi.  16,  fig.  36. 


54  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Miller,  1947:100-101,  pi.  76,  figs.  2-3;  pi.  77,  figs.  1-2; 

pi.  92,  figs.  6-7. 

Primary  Type  Material.  UCMP  holotype  31089,  Ardath 
Shale,  UCMP  locality  3989. 

Molluscan  Stage  Range.  “Capay”  through  “Domengine.” 

Geographic  Distribution.  San  Diego  through  central  Cal- 
ifornia. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  374,  458. 

Remarks.  Specimens  were  found  only  in  the  “Stewart  bed.” 
The  best  preserved  specimens  are  from  locality  374  where 
one  juvenile  (Fig.  13a)  and  an  adult  specimen  were  found. 
As  reported  by  Schenck  (1931),  a hypotype  of  A.  myrlae  is 
from  UCMP  7004  = CSUN  locality  374. 

Subclass  Coleoidea 
Order  Sepiida 
Family  Indeterminate 

Spirulimorph  Sepiid 

Figure  13b 

Spirulimorph  sepiid  Squires,  1983a:360,  figs.  2j-k. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  493. 

Remarks.  Only  a single  partial  phragmocone  and  associ- 
ated external  mold  were  found  in  the  Llajas  Formation.  In 
the  absence  of  the  rostrum  and  the  apical  part  of  the  phrag- 
mocone, the  specimen  and  mold  are  unidentifiable.  This  is 
the  first  sepiid  in  the  Eocene  of  western  North  America 
(Squires,  1983a). 


Phylum  Arthropoda 

Class  Malacostraca 

Subclass  Eumalacostraca 

Order  Decapoda 

Suborder  Pleocyemata 

Infraorder  Brachyura 

Section  Brachyrhyncha 

Superfamily  Xanthoidea 

Family  Goneplacidae  MacLeay,  1838 

Subfamily  Carcinoplacinae 
Milne-Edwards,  1852 

Genus  Glyphithyreus  Reuss,  1859 

Type  Species.  By  original  designation,  Glyphithyreus  for- 
mosus  Reuss,  1859  ( = Plagioloph us  wetherelli  Bell,  1858). 


28 


Location  of  measured  section 

Contact 

Fault 

Generalized  attitude  of  beds 
Section  number 
Paved  road 

Improved  secondary  road 
Unimproved  secondary  road 


|Tsp|  Sespe  Format 
[ T1  | Llajas  Format 
P US  Shallow-mar 

□ 


ne  (regressive)  facies 

Outer  shelf  & slope  facies  with 
localized  turbidite-filled  channels 

Shallow-marine  (transgressive)  facies 

Coastal  alluvial-fan  facies 

Santa  Susana  Formation 


Figure  14.  Index  map  to  the  Llajas  Formation  showing  locations 
of  areas  used  as  fossil-collecting  locality  maps  in  Figures  15-19.  An 
explanation  of  symbols  used  on  the  locality  maps  is  also  given. 


Glyphithyreus  weaveri  (Rathbun,  1926) 

Figure  13c 

Cancer ? sp.  Weaver,  1905:123,  pi.  13,  fig.  11. 

Cancer  (?)  sp.  A Dickerson,  1 9 1 6:pl.  42,  fig.  11. 
Plagiolophus  weaveri  Rathbun,  1926:35-37,  pi.  9,  figs.  5-6. 
Orr  and  Kooser,  1971:157,  text  figure  3a,  figs.  4a-c,  figs. 
5a-i. 

Primary  Type  Material.  USNM  holotype  353351,  Eocene 
strata.  Salt  Creek,  5.2  km  north-northeast  of  Joaquin  Rocks, 
south  part  of  NW  Vi  of  section  15,  T 18  S,  R 14  E,  Fresno 
County,  California. 

Molluscan  Stage  Range.  “Capay”  through  “Domengine.” 
Geographic  Distribution.  San  Diego,  California  through 
southwestern  Oregon. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  471b,  548. 

Remarks.  Although  fragments  of  brachyuran  chelipeds  were 
found  at  widely  scattered  localities  in  the  Llajas  Formation, 
carapaces  were  found  only  at  localities  471b  and  548.  At 
locality  548,  most  of  the  specimens  have  also  one  or  both 
chelipeds  intact. 

Phylum  Echinodermata 
Subphylum  Echinozoa 
Class  Echinoidea 
Subclass  Euechinoidea 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  55 


• 462 


. 476 

• 475 

• 489 


. 541 
. 473 

. 472,458, 459, 

n477,  478  , 
479, 480 


.481 
•440 
. 439 
* 537 
•453 


• 442 

• 441 


I ® 


. 452,705 

• 454,706 

• 542 


Figure  15.  Geologic  map  showing  CSUN  fossil-collecting  localities,  Llajas  Formation,  type  section  area.  Accompanying  columnar  section 
shows  stratigraphic  position  of  the  fossil-collecting  localities.  See  Figure  14  for  explanation  of  symbols. 


Superorder  Atelostomata 

Order  Spatangoida 

Suborder  Hemiasterina 

Family  Schizasteridae  Lambert,  1902 

Genus  Schizaster  L.  Agassiz,  1836 

Type  Species.  By  subsequent  designation  (ICZN,  1954, 
opin.  209),  Schizaster  studeri  L.  Agassiz,  1836. 

Schizaster  diabloensis  Kew,  1920 

Figure  1 3d 

Schizaster  diabloensis  Kew,  1920:150-151,  pi.  41,  figs.  5a- 
c.  Clark  and  Woodford,  1927:123,  pi.  22,  fig.  14.  Clark, 
1929:pl.  4,  fig.  13.  Grant  and  Hertlein,  1938:120. 

Primary  Type  Material.  UCMP  holotype  1 1387,  Eocene 
strata,  UCMP  locality  1427. 

Molluscan  Stage  Range.  Upper  Paleocene  through  “Do- 
mengine.” 

Geographic  Distribution.  Simi  Valley  through  Marysville 
Buttes,  California. 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  374,  458,  478.  Outer  shelf  and  slope  facies: 
CSUN  locality  482.  Shallow-marine  (regressive)  facies:  CSUN 
locality  547. 


Remarks.  Preservation  is  as  internal  molds  and  most  spec- 
imens are  somewhat  crushed.  At  locality  547,  5.  diabloensis 
is  the  only  megafossil  present. 

Clark  and  Woodford  (1927)  reported  this  species  from  the 
type  section  area  of  the  Meganos  Formation.  Marincovich 
(1977:251)  considered  the  formation  to  be  late  Paleocene  in 
age. 

Phylum  Chordata 

Class  Chondrichthyes 

Subclass  Elasmobranchii 

Order  Lamniformes 

Family  Odontaspididae 
Muller  and  Henle,  1837 

Genus  Odontaspis  L.  Agassiz,  1838 

Type  Species.  By  monotypy,  Odontaspis ferox  Risso,  1810. 

Odontaspis  sp. 

Figure  13e 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  localities  374,  445. 


56  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Figure  16.  Geologic  map  showing  CSUN  fossil-collecting  localities,  Llajas  Formation,  lower  Las  Llajas  Canyon  area.  Accompanying  columnar 
section  shows  stratigraphic  position  of  the  fossil-collecting  localities.  See  Figure  14  for  explanation  of  symbols. 


Remarks.  Only  a few  specimens  were  found  in  the  Llajas 
Formation. 

Family  Lamnidae  Muller  and  Henle,  1838 
Genus  Isurus  Rafinesque,  1 809 

Type  Species.  By  original  designation,  Isurus  oxyrhynchus 
Rafinesque,  1809. 

Isurus  cf.  I.  praecursor  (Leriche,  1906) 

Figure  13f 

Local  Occurrence.  Shallow-marine  (transgressive)  facies: 
CSUN  locality  371. 

Remarks.  Only  one  specimen  was  found  in  the  Llajas  For- 
mation. 

LOCALITIES 

CSUN  fossil-collecting  localities  made  by  the  author  in  the 
course  of  this  study  are  listed  first.  Localities  of  other  insti- 
tutions mentioned  in  this  report  follow  in  alphabetical  order. 

All  CSUN  localities  are  in  the  Llajas  Formation,  Simi 
Valley  area,  California.  Unless  otherwise  noted,  they  are  in 
the  United  States  Geological  Survey  7.5-minute  topographic 


quadrangle  of  Santa  Susana,  California  (195 1),  photorevised 
1969.  In  some  cases,  a locality  is  in  the  Santa  Susana  quad- 
rangle, but  the  section  corner  used  for  referencing  is  in  the 
United  States  Geological  Survey  7.5-minute  topographic 
quadrangle  of  Oat  Mountain,  California  (1952),  photorevised 
1969.  These  localities  are  so  noted.  Abbreviations  used  are 
United  States  Geological  Survey  (USGS),  feet  (ft.),  meters 
(m),  township  (T),  range  (R),  north  (N),  south  (S),  east  (E), 
and  west  (W).  Distances  are  given  in  both  English  and  metric 
units,  but  map  contour  elevations  are  given  in  English  units 
only. 

CSUN  localities  equivalent  to  localities  of  University  of 
California  Museum  of  Paleontology  (Berkeley)  (UCMP); 
University  of  California,  Los  Angeles  (UCLA);  and  Califor- 
nia Institute  of  Technology  (CIT)  are  so  indicated.  CSUN 
localities  are  denoted  also  as  to  which  marine  facies  (Squires, 
1981)  they  belong.  The  general  location  and  the  relative 
stratigraphic  position  of  each  CSUN  locality  are  shown  in 
Figures  14  through  19. 

CSUN  LOCALITIES 

371.  At  elevation  of  2000  ft.  on  south  side  of  a side  canyon 
to  Devil  Canyon,  1275  ft.  (389  m)  south  and  1150  ft.  (350 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  57 


1 km 

N 


BASE  MAP  FROM  SANTA  SUSANA  SUSANA  (1951)  AND  OAT  MOUNTAIN  (1952).  CA., 
7.5-MINUTE  QUADRANGLES,  BOTH  PHOTOREVISED  1969. 


Figure  17.  Geologic  map  showing  CSUN  fossil-collecting  localities,  Llajas  Formation,  Devil  Canyon  area.  Accompanying  columnar  section 
shows  stratigraphic  position  of  the  fossil-collecting  localities.  See  Figure  14  for  explanation  of  symbols. 


m)  west  of  NE  comer  of  section  26,  T 3 N,  R 1 7 W.  Locality 
is  in  the  Santa  Susana  quadrangle,  but  the  section  comer  is 
in  the  Oat  Mountain  quadrangle.  [Shallow-marine  (trans- 
gressive) facies.] 

372.  At  elevation  of  2035  ft.  on  west  side  of  stream  bank 
of  Devil  Canyon,  800  ft.  (244  m)  south  and  1200  ft.  (366  m) 
west  of  NE  comer  of  section  26,  T 3 N,  R 17  W.  Locality  is 
in  the  Santa  Susana  quadrangle,  but  the  section  comer  is  in 
the  Oat  Mountain  quadrangle.  [Shallow-marine  (transgres- 
sive) facies.] 

373.  At  elevation  of  1900  ft.  on  east  side  of  stream  bank 
of  Devil  Canyon,  450  ft.  (137  m)  south  and  850  ft.  (259  m) 
west  of  NE  comer  of  section  26,  T 3 N,  R 17  W.  Locality  is 
in  the  Santa  Susana  quadrangle,  but  the  section  corner  is  in 
the  Oat  Mountain  quadrangle.  [Shallow-marine  (transgres- 
sive) facies.] 

374.  At  elevation  of  1700  ft.  on  a small  cliff  on  south  side 
of  a side  canyon  to  Las  Llajas  Canyon,  1950  ft.  (594  m)  north 
and  1825  ft.  (556  m)  east  of  SE  comer  of  section  29,  T 3 N, 
R 17  W.  Locality  is  in  the  “Stewart  bed”  and  is  equivalent 
to  UCMP  locality  7004,  UCLA  locality  2312,  and  CIT  lo- 
cality 206.  [Shallow-marine  (transgressive)  facies.] 

438.  At  elevation  of  1800  ft.  on  east  side  of  stream  bank 


of  Las  Llajas  Canyon,  7000  ft.  (2 1 34  m)  N86°E  of  NE  comer 
of  section  29,  T 3 N,  R 1 7 W.  Locality  is  equivalent  to  UCLA 
locality  2775.  [Shallow-marine  (transgressive)  facies.] 

439.  At  elevation  of  1425  ft.  on  a ridge,  800  ft.  (244  m) 
south  and  1900  ft.  (579  m)  east  of  NW  corner  of  section  31, 
T 3 N,  R 1 7 W.  [Shallow-marine  (transgressive)  facies.] 

440.  At  elevation  of  1475  ft.  on  a ridge,  600  ft.  (183  m) 
south  and  1875  ft.  (571  m)  east  of  NW  comer  of  section  31, 
T 3 N,  R 17  W.  [Shallow-marine  (transgressive)  facies.] 

441 . At  elevation  of  1 1 60  ft.  on  west  side  of  Chivo  Canyon 
near  its  mouth,  1890  ft.  (576  m)  south  and  400  ft.  (122  m) 
west  of  NE  comer  of  section  31,  T 3 N,  R 17  W.  [Shallow- 
marine  (transgressive)  facies.] 

442.  At  elevation  of  1 1 85  ft.  on  west  side  of  Chivo  Canyon 
near  its  mouth,  1600  ft.  (488  m)  south  and  510  ft.  (155  m) 
west  of  NE  comer  of  section  31,  T 3 N,  R 17  W.  [Shallow- 
marine  (transgressive)  facies.] 

443.  At  elevation  of  1 230  ft.  on  east  side  of  Chivo  Canyon 
near  its  mouth,  1370  ft.  (418  m)  south  and  200  ft.  (61  m) 
west  of  NE  comer  of  section  31,  T 3 N,  R 17  W.  [Shallow- 
marine  (transgressive)  facies.] 

444.  At  elevation  of  1 585  ft.  on  a small  cliff  on  north  side 
of  a side  canyon  to  Las  Llajas  Canyon,  2500  ft.  (762  m)  north 


58  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


468b,  516 

• 438,468a,  515,  525 

514  . 513 

• 467  ’ 524 

• 523 
’ 521 

. 518 

• 520 


. 460 
• 517 


• 466 


• 465 

• 463 


Figure  18.  Geologic  map  showing  CSUN  fossil-collecting  localities,  Llajas  Formation,  upper  Las  Llajas  Canyon  area.  Accompanying  columnar 
section  shows  stratigraphic  position  of  the  fossil-collecting  localities.  See  Figure  14  for  explanation  of  symbols. 


and  1375  ft.  (419  m)  east  of  SE  comer  of  section  29,  T 3 N, 
R 17  W.  Locality  is  in  the  “Stewart  bed.”  [Shallow-marine 
(transgressive)  facies.] 

445.  At  elevation  of  1485  ft.  on  a small  cliff  on  west  side 
of  Las  Llajas  Canyon,  2820  ft.  (859  m)  north  and  850  ft.  (259 
m)  east  of  SE  comer  of  section  29,  T 3 N,  R 1 7 W.  Locality 
is  in  the  “Stewart  bed.”  [Shallow-marine  (transgressive)  fa- 
cies.] 

446.  At  elevation  of  1 760  ft.  in  roadcut  along  ridge  on  west 
side  of  Las  Llajas  Canyon,  3460  ft.  (1055  m)  north  and  200 
ft.  (61  m)  east  of  SE  corner  of  section  29,  T 3 N,  R 17  W. 
[Outer  shelf  and  slope  facies.] 

447.  At  elevation  of  1550  ft.  on  a small  cliff  on  west  side 
of  Las  Llajas  Canyon,  2780  ft.  (847  m)  north  and  550  ft.  (168 
m)  east  of  SE  comer  of  section  29,  T 3 N,  R 17  W.  Locality 
is  in  the  “Stewart  bed.”  [Shallow-marine  (transgressive)  fa- 
cies.] 

449.  At  elevation  of  1700  ft.  on  a small  cliff  on  west  side 
of  Las  Llajas  Canyon,  1825  ft.  (556  m)  north  and  200  ft.  (61 
m)  west  of  SE  comer  of  section  29,  T 3 N,  R 17  W.  Locality 
is  in  the  “Stewart  bed.”  [Shallow-marine  (transgressive)  fa- 
cies.] 

450.  At  elevation  of  1600  ft.  on  a small  cliff  on  west  side 
of  Las  Laljas  Canyon,  2450  ft.  (747  m)  north  and  125  ft.  (38 
m)  east  of  SE  comer  of  section  29,  T 3 N,  R 17  W.  Locality 


is  in  the  “Stewart  bed.”  [Shallow-marine  (transgressive)  fa- 
cies.] 

451.  At  elevation  of  1715  ft.  on  a small  cliff  on  a hillside, 
2100  ft.  (640  m)  north  and  2100  ft.  (640  m)  west  of  SE  corner 
of  section  29,  T 3 N,  R 17  W.  Locality  is  in  the  “Stewart 
bed.”  [Shallow-marine  (transgressive)  facies  ] 

452.  At  elevation  of  1 285  ft.  on  west  side  of  a small  canyon 
branching  north  near  mouth  of  Las  Llajas  Canyon,  1650  ft. 
(503  m)  south  and  550  ft.  (168  m)  east  of  NW  corner  of 
section  32,  T 3 N,  R 1 7 W.  [Interfingering  coastal  alluvial- 
fan  facies  and  shallow-marine  (transgressive)  facies.] 

453.  At  elevation  of  1 275  ft.  on  west  side  of  Chivo  Canyon, 
800  ft.  (243  m)  south  and  700  ft.  (213  m)  west  of  NE  comer 
of  section  3 1 , T 3 N,  R 1 7 W.  [Shallow-marine  (transgressive) 
facies.] 

454.  At  elevation  of  1 3 1 0 ft.  on  west  side  of  a small  canyon 
branching  north  near  mouth  of  Las  Llajas  Canyon,  1 100  ft. 
(335  m)  south  and  810  ft.  (247  m)  east  of  NW  comer  of 
section  32,  T 3 N,  R 17  W.  [Interfingermg  coastal  alluvial- 
fan  facies  and  shallow-marine  (transgressive)  facies.] 

455.  At  elevation  of  1475  ft.  on  east  side  of  stream  bank 
of  Las  Llajas  Canyon,  2500  ft.  (762  m)  north  and  625  ft.  ( 191 
m)  east  of  SE  comer  of  section  29,  T 3 N,  R 17W.  [Shallow- 
marine  (transgressive)  facies.] 

457.  At  elevation  of  1 300  ft.  on  east  side  of  Chivo  Canyon, 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  59 


700  m 


TsP _/ 

■ L X 

-V 

\ 

1 

1 

• 

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544 

Tl 

19 

cv'L- 

// 

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CO 

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Tsp 


• 544 


• 547 


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eSEEeje*-, 


548 

543 

704 


. 546 
,703  . . 

! 702 
'700,  701 


BASE  MAP  FROM  SANTA  SUSANA  (1951,  PHOTOREVISED  1969)  AND  THOUSAND  OAKS  (1950,  PHOTOREVISED  1967),  CA..  7.5-MINUTE  OUADRANGLES. 

Figure  19.  Geologic  maps  showing  CSUN  fossil-collecting  localities,  Llajas  Formation,  at  various  areas  along  the  south  side  of  Simi  Valley. 
Accompanying  columnar  sections  shows  stratigraphic  position  of  the  fossil-collecting  localities.  See  Figure  14  for  explanation  of  symbols,  a. 
Long  Canyon  area.  b.  Lower  Bus  Canyon  area.  c.  Simi  Arroyo-Peppertree  Lane  area. 


900  ft.  (274  m)  north  and  300  ft.  (91  m)  west  of  SE  comer 
of  section  30,  T 3 N,  R 17  W.  Bed  is  the  same  one  exposed 
at  CSUN  localities  4691  and  511.  [Shallow-marine  (trans- 
gressive) facies.] 

458.  At  elevation  of  1300  ft.  along  south  side  of  a ranch 
road  that  leads  up  south  side  of  Chivo  Canyon,  1550  ft.  (472 
m)  north  and  400  ft.  (122  m)  east  of  SW  comer  of  section 
29,  T 3 N,  R 1 7 W.  Locality  is  in  the  “Stewart  bed."  [Shallow- 
marine  (transgressive)  facies.] 

459.  At  elevation  of  1400  ft.  on  north  side  of  Chivo  Can- 
yon, 1500  ft.  (457  m)  north  and  400  ft.  (122  m)  west  of  SE 
comer  of  section  30,  T 3 N,  R 17  W.  Locality  is  in  the 
“Stewart  bed.”  [Shallow-marine  (transgressive)  facies.] 

460.  At  elevation  of  2150  ft.  on  a hillside  just  north  of  a 
ranch  road,  6700  ft.  (2042  m)  N67°E  of  NE  corner  of  section 
29,  T 3 N,  R 17  W.  [Shallow-marine  (transgressive)  facies.] 

462.  At  elevation  of  1225  ft.  on  a hillside  just  north  of  an 
oilfield  road,  2250  ft.  (686  m)  south  and  2675  ft.  (815  m) 
west  of  NE  comer  of  section  36,  T 3 N,  R 18  W.  Locality  is 
near  the  Llajas-Sespe  contact.  [Shallow-marine  (regressive) 
facies.] 


463.  At  elevation  of  1650  ft.  in  streambed.  Las  Llajas 
Canyon,  3725  ft.  (1 1 35  m)  N89°E  of  SE  comer  of  section  29, 
T 3 N,  R 17  W.  [Shallow-marine  (transgressive)  facies.] 

465.  At  elevation  of  2100  ft.  on  south  side  of  a hill,  5480 
ft.  (1670  m)  N70°E  of  NE  comer  of  section  29,  T 3 N,  R 1 7 
W.  [Shallow-marine  (transgressive)  facies.] 

466.  At  elevation  of  2 1 50  ft.  along  east  side  of  a hill,  5700 
ft.  (1738  m)  N69°E  of  NE  comer  of  section  29,  T 3 N,  R 17 
W.  [Shallow-marine  (transgressive)  facies.] 

467.  At  elevation  of  1 900  ft.  on  west  side  of  stream  bank 
of  El  Toro  Canyon,  7900  ft.  (2408  m)  N69°W  of  SE  comer 
of  section  23,  T 3 N,  R 1 7 W.  Locality  is  in  the  Santa  Susana 
quadrangle,  but  the  section  comer  is  in  the  Oat  Mountain 
quadrangle.  [Shallow-marine  (transgressive)  facies.] 

468a.  At  elevation  of  1975  ft.  on  north  side  of  stream  bank 
of  Las  Llajas  Canyon,  6675  ft.  (2034  m)  N69°W  of  SE  comer 
of  section  23,  T 3 N,  R 1 7 W.  Locality  is  in  the  Santa  Susana 
quadrangle,  but  the  section  comer  is  in  the  Oat  Mountain 
quadrangle.  Bed  is  the  same  one  exposed  at  CSUN  locality 
515.  [Shallow-marine  (transgressive)  facies.] 

468b.  Bed  1 0 ft.  (3  m)  stratigraphically  above  locality  468a. 


60  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Bed  is  the  same  one  exposed  at  CSUN  locality  516.  [Shallow- 
marine  (transgressive)  facies.] 

469a.  At  elevation  of  1400  ft.  on  north  side  of  a small 
canyon  branching  west  from  Chivo  Canyon,  100  ft.  (31  m) 
north  and  1650  ft.  (503  m)  west  of  SE  comer  of  section  30, 
T 3 N,  R 17  W.  Locality  is  at  the  type  section  of  the  Llajas 
Formation,  144  ft.  (44  m)  above  the  base  of  the  formation. 
[Interfingering  coastal  alluvial-fan  facies  and  shallow-marine 
(transgressive)  facies.] 

469b.  Bed  26  ft.  (8  m)  stratigraphically  above  locality  469a, 
on  a traverse  bearing  N10°W  from  locality  469a.  [Interfin- 
gering coastal  alluvial-fan  facies  and  shallow-marine  (trans- 
gressive) facies.] 

469c.  Bed  1 3 ft.  (4  m)  stratigraphically  above  locality  469b, 
on  a traverse  bearing  N10°W  from  locality  469b.  [Interfin- 
gering coastal  alluvial-fan  facies  and  shallow-marine  (trans- 
gressive) facies.] 

469d.  Bed  1 3 ft.  (4  m)  stratigraphically  above  locality  469c, 
on  a traverse  bearing  N10°W  from  locality  469c.  [Interfin- 
gering coastal  alluvial-fan  facies  and  shallow-marine  (trans- 
gressive) facies.] 

469e.  Bed  20  ft.  (6  m)  stratigraphically  above  locality  469d, 
on  a traverse  bearing  N10°W  from  locality  469d.  [Shallow- 
marine  (transgressive)  facies.] 

469f.  Bed  6.5  ft.  (2  m)  stratigraphically  above  locality  469e, 
on  a traverse  bearing  N10°W  from  locality  469e.  [Shallow- 
marine  (transgressive)  facies.] 

469g.  Bed  6.5  ft.  (2  m)  stratigraphically  above  locality  469f, 
on  a traverse  bearing  N10°W  from  locality  469f.  [Shallow- 
marine  (transgressive)  facies.] 

469h.  Bed  1 6 ft.  (5  m)  stratigraphically  above  locality  469g, 
on  a traverse  bearing  N10°W  from  locality  469g.  [Shallow- 
marine  (transgressive)  facies.] 

4691.  Bed  9 ft.  (3  m)  stratigraphically  above  locality  469h, 
on  a traverse  bearing  N10°W  from  locality  469h.  [Shallow- 
marine  (transgressive)  facies.] 

469j.  Bed  75  ft.  (19  m)  stratigraphically  above  locality 
469g,  on  a traverse  bearing  N10°W  from  locality  469g.  [Shal- 
low-marine (transgressive)  facies.] 

469k.  Bed  20  ft.  (6  m)  stratigraphically  above  locality  469g, 
on  a traverse  bearing  N10°W  from  locality  469j.  [Shallow- 
marine  (transgressive)  facies.] 

4691.  Bed  13  ft.  (4  m)  stratigraphically  above  locality  469k, 
on  a traverse  bearing  N55°W  from  locality  469k.  Bed  is  the 
same  one  exposed  at  CSUN  localities  457  and  511.  [Shallow- 
marine  (transgressive)  facies.] 

470a.  At  elevation  of  1550  ft.  on  south  side  of  an  amphi- 
theater on  west  side  of  Chivo  Canyon,  500  ft.  (152  m)  north 
and  2100  ft.  (640  m)  west  of  SE  comer  of  section  30,  T 3 N, 
R 17  W.  Locality  is  at  the  type  section  of  the  Llajas  For- 
mation, 266  ft.  (81  m)  stratigraphically  above  locality  4691. 
[Shallow-marine  (transgressive)  facies.] 

470b.  Bed  62  ft.  (19  m)  stratigraphically  above  locality 
470a,  on  a traverse  bearing  N60°W  from  locality  470a.  [Shal- 
low-marine (transgressive)  facies.] 

470c.  Bed  33  ft.  (10  m)  stratigraphically  above  locality 
470b,  on  a traverse  bearing  N60°W  from  locality  470b.  [Shal- 
low-marine (transgressive)  facies.] 


471a.  At  elevation  of  1450  ft.  on  west  side  of  an  amphi- 
theater on  west  side  of  Chivo  Canyon,  600  ft.  (183  m)  north 
and  2125  ft.  (655  m)  west  of  SE  comer  of  section  30,  T 3 N, 
R 17  W.  Locality  is  at  the  type  section  of  the  Llajas  For- 
mation, 125  ft.  (38  m)  stratigraphically  above  locality  470c. 
[Shallow-marine  (transgressive)  facies.] 

471b.  Bed  144  ft.  (44  m)  stratigraphically  above  locality 
471a,  on  a traverse  bearing  N27°W  from  locality  471a.  [Shal- 
low-marine (transgressive)  facies.] 

471c.  Bed  20  ft.  (6  m)  stratigraphically  above  locality  471b, 
on  a traverse  bearing  N27°W  from  locality  471b.  [Shallow- 
marine  (transgressive)  facies.] 

47!d.  Bed  39  ft.  (12  m)  stratigraphically  above  locality 
47  lc,  on  a traverse  bearing  N27°W  from  locality  471c.  [Shal- 
low-marine (transgressive)  facies.] 

471e.  Bed  79  ft.  (24  m)  stratigraphically  above  locality 
47  Id,  on  a traverse  bearing  N27°W  from  locality  47  Id.  [Shal- 
low-marine (transgressive)  facies.] 

472.  At  elevation  of  1600  ft.  on  a ridge,  1000  ft.  (305  m) 
north  and  2300  ft.  (701  m)  west  of  SE  corner  of  section  30, 
T 3 N,  R 1 7 W.  Locality  is  at  the  type  section  of  the  Llajas 
Formation  and  in  the  “Stewart  bed,”  46  ft.  (14  m)  strati- 
graphically above  locality  47  le.  [Shallow-marine  (transgres- 
sive) facies.] 

473.  At  elevation  of  1650  ft.  on  a ridge,  1050  ft.  (320  m) 
north  and  2300  ft.  (701  m)  west  of  SE  corner  of  section  30, 
T 3 N,  R 1 7 W.  Locality  is  at  the  type  section  of  the  Llajas 
Formation,  46  ft.  (14  m)  stratigraphically  above  locality  472. 
The  bed  that  this  locality  occurs  in  probably  represents  a 
shallow-marine  (transgressive)  facies  bed  that  interfingers  with 
the  outer  shelf  and  slope  facies. 

475.  At  elevation  of  1625  ft.  on  a ridge,  1300  ft.  (396  m) 
north  and  3100  ft.  (945  m)  west  of  SE  corner  of  section  30, 
T 3 N,  R 1 7 W.  Locality  is  at  the  type  section  of  the  Llajas 
Formation,  approximately  446  ft.  (136  m)  stratigraphically 
above  locality  473.  [Shallow-marine  (regressive)  facies.] 

476.  Bed  13  ft.  (4  m)  stratigraphically  above  locality  475, 
on  a traverse  bearing  N15°W  from  locality  475.  [Shallow- 
marine  (regressive)  facies.] 

477.  At  elevation  of  1625  ft.  on  a ridge,  1200  ft.  (366  m) 
north  and  1700  ft.  (518  m)  west  of  SE  comer  of  section  30, 
T 3 N,  R 17  W.  Locality  is  in  the  “Stewart  bed.”  [Shallow- 
marine  (transgressive)  facies.] 

478.  At  elevation  of  1735  ft.  just  below  a saddle  along  a 
ridge,  680  ft.  (270  m)  north  and  2670  ft.  (814  m)  west  of  SE 
comer  of  section  30,  T 3 N,  R 1 7 W.  Locality  is  in  the 
“Stewart  bed.”  [Shallow-marine  (transgressive)  facies.] 

479.  At  elevation  of  1775  ft.  just  below  a saddle  along  a 
ridge,  650  ft.  (198  m)  north  and  2700  ft.  (823  m)  west  of  SE 
comer  of  section  30,  T 3 N,  R 17  W.  Locality  is  in  the 
“Stewart  bed.”  [Shallow-marine  (transgressive)  facies.] 

480.  At  elevation  of  1400  ft.  near  a canyon  bottom,  200 
ft.  (61  m)  north  and  3400  ft.  (1036  m)  west  of  SE  comer  of 
section  30,  T 3 N,  R 17  W.  Locality  is  in  the  “Stewart  bed.” 
[Shallow-marine  (transgressive)  facies.] 

481.  At  elevation  of  1500  ft.  on  a ridge,  325  ft.  (99  m) 
south  and  1800  ft.  (549  m)  east  of  NE  comer  of  section  31, 
T 3 N,  R 1 7 W.  [Shallow-marine  (transgressive)  facies.] 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  61 


482.  At  elevation  of  1435  ft.  in  a roadcut,  1 100  ft.  (335 
m)  south  and  300  ft.  (91  m)  west  of  NE  corner  of  section  36, 
T 3 N,  R 18  W.  [Outer  shelf  and  slope  facies.] 

483.  At  elevation  of  1525  ft.  on  west  side  of  a ridge,  925 
ft.  (282  m)  south  and  175  ft.  (53  m)  west  of  NE  comer  of 
section  36,  T 3 N,  R 18  W.  [Outer  shelf  and  slope  facies.] 

484.  At  elevation  of  2340  ft.  on  a ridge,  6000  ft.  (1829  m) 
N68°W  of  SE  comer  of  section  26,  T 3 N,  R 17  W.  Locality 
is  in  the  Santa  Susana  quadrangle,  but  the  section  corner  is 
in  the  Oat  Mountain  quadrangle.  [Shallow-marine  (trans- 
gressive) facies.] 

485.  Bed  1 35  ft.  (4 1 m)  stratigraphically  below  locality  484, 
on  a traverse  bearing  S23°E  from  locality  484.  [Shallow- 
marine  (transgressive)  facies.] 

486.  Bed  69  ft.  (21  m)  stratigraphically  below  locality  485, 
on  a traverse  bearing  S23°E  from  locality  485.  [Shallow- 
marine  (transgressive)  facies.] 

487.  Bed  1 74  ft.  (53  m)  stratigraphically  below  locality  486, 
on  a traverse  bearing  S23°E  from  locality  486.  [Shallow- 
marine  (transgressive)  facies.] 

488.  At  elevation  of  2175  ft.  on  a ridge,  5650  ft.  (1722  m) 
N64°W  of  SE  comer  of  section  26,  T 3 N,  R 17  W.  Locality 
is  in  the  Santa  Susana  quadrangle,  but  the  section  comer  is 
in  the  Oat  Mountain  quadrangle.  Locality  is  in  the  “Stewart 
bed.”  [Shallow-marine  (transgressive)  facies.] 

489.  At  elevation  of  1250  ft.  on  east  side  of  a canyon,  125 
ft.  (38  m)  south  of  NW  corner  of  section  31,  T 3 N,  R 17 
W.  [Shallow-marine  (regressive)  facies.] 

490.  At  elevation  of  1900  ft.  in  a saddle  along  a ridge,  600 
ft.  (183  m)  north  and  800  ft.  (244  m)  east  of  SE  corner  of 
section  29,  T 3 N,  R 17  W.  [Shallow-marine  (transgressive) 
facies.] 

491.  At  elevation  of  1 840  ft.  on  east  side  of  a small  canyon, 
500  ft.  (145  m)  north  and  1 100  ft.  (320  m)  east  of  SE  comer 
of  section  29,  T 3 N,  R 1 7 W.  [Shallow-marine  (transgressive) 
facies.] 

492.  At  elevation  of  2230  ft.  in  a saddle  along  a ridge,  1 700 
ft.  (518  m)  north  and  3225  ft.  (983  m)  east  of  SE  comer  of 
section  29,  T 3 N,  R 1 7 W.  [Shallow-marine  (transgressive) 
facies.] 

493.  At  elevation  of  2225  ft.  in  saddle  along  a ridge,  1825 
ft.  (556  m)  north  and  3450  ft.  (1052  m)  east  of  SE  comer  of 
section  29,  T 3 N,  R 17  W.  Locality  is  in  the  “Stewart  bed.” 
[Shallow-marine  (transgressive)  facies.] 

494.  At  elevation  of  2050  ft.  on  a small  cliff,  5350  ft.  (1630 
m)  N62°W  of  SE  comer  of  section  26,  T 3 N,  R 1 7 W.  Locality 
is  in  the  Santa  Susana  quadrangle,  but  the  section  corner  is 
in  the  Oat  Mountain  quadrangle.  Locality  is  in  the  “Stewart 
bed.”  [Shallow-marine  (transgressive)  facies.] 

495.  At  elevation  of  2050  ft.  on  a small  cliff,  4725  ft.  (1440 
m)  N60°W  of  SE  comer  of  section  26,  T 3 N,  R 1 7 W.  Locality 
is  in  the  Santa  Susana  quadrangle,  but  the  section  comer  is 
in  the  Oat  Mountain  quadrangle.  Locality  is  in  the  “Stewart 
bed.”  [Shallow-marine  (transgressive)  facies.] 

496.  At  elevation  of  2050  ft.  on  a hillside,  4100  ft.  (1250 
m)N59°W  ofSE  comer  of  section  26,  T 3 N,  R 17  W.  Locality 
is  in  the  Santa  Susana  quadrangle,  but  the  section  comer  is 


in  the  Oat  Mountain  quadrangle.  Locality  is  in  the  “Stewart 
bed.”  [Shallow-marine  (transgressive)  facies.] 

497.  At  elevation  of  2185  ft.  on  south  side  of  ridge,  3685 
ft.  (1 123  m)  N58°W  of  SE  comer  of  section  26,  T 3 N,  R 17 
W.  Locality  is  in  the  Santa  Susana  quadrangle,  but  the  section 
comer  is  in  the  Oat  Mountain  quadrangle.  Locality  is  in  the 
“Stewart  bed.”  [Shallow-marine  (transgressive)  facies.] 

498.  At  elevation  of  1850  ft.  on  west  side  of  stream  bank 
of  Devil  Canyon,  1550  ft.  (472  m)  south  and  600  ft.  (183  m) 
west  of  NE  comer  of  section  26,  T 3 N,  R 1 7 W of  the  USGS 

7.5-minute  topographic  quadrangle  of  Oat  Mountain,  Cali- 
fornia (1952),  photorevised  1969.  [Shallow-marine  (trans- 
gressive) facies.] 

499.  At  elevation  of  1850  ft.  on  west  side  of  stream  bank 
of  Devil  Canyon,  1400  ft.  (427  m)  south  and  630  ft.  (192  m) 
west  of  NE  comer  of  section  26,  T 3 N,  R 17  W of  the  USGS 

7.5-minute  topographic  quadrangle  of  Oat  Mountain,  Cali- 
fornia (1952),  photorevised  1969.  [Shallow-marine  (trans- 
gressive) facies.] 

500.  At  elevation  of  1875  ft.  on  east  side  of  stream  bank 
of  Devil  Canyon,  800  ft.  (244  m)  south  and  675  ft.  (206  m) 
west  of  NE  comer  of  section  26,  T 3 N,  R 1 7 W of  the  USGS 

7.5-minute  topographic  quadrangle  of  Oat  Mountain,  Cali- 
fornia (1952),  photorevised  1969.  [Shallow-marine  (trans- 
gressive) facies.] 

501.  At  elevation  of  1500  ft.  on  east  side  of  a small  canyon 
branching  northwest  of  Las  Llajas  Canyon,  1400  ft.  (427  m) 
north  and  1400  ft.  (427  m)  west  of  SE  comer  of  section  29, 
T 3 N,  R 1 7 W.  [Interfingering  coastal  alluvial-fan  facies  and 
shallow-marine  (transgressive)  facies.] 

502.  At  elevation  of  2150  ft.  on  a ridge,  5700  ft.  (1737  m) 
N70°W  of  SE  comer  of  section  26,  T 3 N,  R 17  W.  Locality 
is  in  the  Santa  Susana  quadrangle,  but  the  section  comer  is 
in  the  Oat  Mountain  quadrangle.  [Interfingering  coastal  al- 
luvial-fan facies  and  shallow-marine  (transgressive)  facies.] 

503.  At  elevation  of  2160  ft.  on  a ridge,  5725  ft.  (1745  m) 
N69°W  of  SE  comer  of  section  26,  T 3 N,  R 17  W.  Locality 
is  in  the  Santa  Susana  quadrangle,  but  the  section  comer  is 
in  the  Oat  Mountain  quadrangle.  [Interfingering  coastal  al- 
luvial-fan facies  and  shallow-marine  (transgressive)  facies.] 

504.  At  elevation  of  2225  ft.  on  a ridge,  5750  ft.  (1753  m) 
N71°W  of  SE  comer  of  section  26,  T 3 N,  R 17  W.  Locality 
is  in  the  Santa  Susana  quadrangle,  but  the  section  comer  is 
in  the  Oat  Mountain  quadrangle.  [Shallow-marine  (trans- 
gressive) facies.] 

505.  Bed  98  ft.  (30  m)  stratigraphically  above  locality  504, 
on  a traverse  bearing  N23°W  from  locality  504.  [Shallow- 
marine  (transgressive)  facies.] 

506.  At  elevation  of  2280  ft.  on  a ridge,  5600  ft.  (1706  m) 
N65°W  of  SE  comer  of  section  26,  T 3 N,  R 17  W.  Locality 
is  in  the  Santa  Susana  quadrangle,  but  the  section  comer  is 
in  the  Oat  Mountain  quadrangle.  [Shallow-marine  (trans- 
gressive) facies.] 

507.  Bed  33  ft.  (10  m)  stratigraphically  above  locality  506, 
on  a traverse  bearing  N6°E  from  locality  506.  [Shallow-ma- 
rine (transgressive)  facies.] 

508.  Bed  6.5  ft.  (2  m)  stratigraphically  above  locality  507, 


62  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


on  a traverse  bearing  N6°E  from  locality  507.  [Shallow-ma- 
rine (transgressive)  facies.] 

509.  Bed  49  ft.  (1 5 m)  stratigraphically  above  locality  508, 
on  a traverse  bearing  N6°E  from  locality  508.  [Shallow-ma- 
rine (transgressive)  facies.] 

510.  Bed  108  ft.  (33  m)  stratigraphically  above  locality 
509,  on  a traverse  bearing  N6°E  from  locality  509.  [Shallow- 
marine  (transgressive)  facies.] 

511.  At  elevation  of  1225  ft.  on  north  sideofChivo  Canyon 
near  a major  fork  in  Chivo  Canyon,  800  ft.  (244  m)  north 
and  800  ft.  (244  m)  west  of  SE  corner  of  section  30,  T 3 N, 
R 17  W.  Bed  is  the  same  one  exposed  at  CSUN  localities 
457  and  4691.  [Shallow-marine  (transgressive)  facies.] 

512a.  At  elevation  of  1 275  ft.  on  east  side  of  a small  canyon 
branching  north  of  Chivo  Canyon,  2275  ft.  (693  m)  south 
and  525  ft.  (160  m)  east  of  NW  corner  of  section  29,  T 3 N, 
R 17  W.  [Shallow-marine  (regressive)  facies.] 

512b.  Bed  20  ft.  (6  m)  stratigraphically  above  locality  5 1 2a, 
on  a traverse  bearing  N35°W  from  locality  512a.  [Shallow- 
marine  (regressive)  facies.] 

512c.  Bed  16  ft.  (5  m)  stratigraphically  above  locality  512b, 
on  a traverse  bearing  N35°W  from  locality  512b.  [Shallow- 
marine  (regressive)  facies.] 

51 2d.  Bed  30  ft.  (9  m)  stratigraphically  above  locality  512b, 
on  a traverse  bearing  N35°W  from  locality  512b.  [Shallow- 
marine  (regressive)  facies.] 

513.  At  elevation  of  1870  ft.  (570  m)  in  the  streambed  of 
Las  Llajas  Canyon,  6825  ft.  (2080  m)  N71°W  of  SE  corner 
of  section  23,  T 3 N,  R 1 7 W.  Locality  is  in  the  Santa  Susana 
quadrangle,  but  the  section  comer  is  in  the  Oat  Mountain 
quadrangle.  [Shallow-marine  (transgressive)  facies.] 

514.  At  elevation  of  1875  ft.  on  north  side  of  stream  bank 
of  Las  Llajas  Canyon,  6800  ft.  (2073  m)  N71°W  ofSE  corner 
of  section  23,  T 3 N,  R 1 7 W.  Locality  is  in  the  Santa  Susana 
quadrangle,  but  the  section  comer  is  in  the  Oat  Mountain 
quadrangle.  [Shallow-marine  (transgressive)  facies.] 

515.  At  elevation  of  1925  ft.  (587  m)  on  the  north  side  of 
stream  bank  of  Las  Llajas  Canyon,  6750  ft.  (2057  m)  N69°W 
of  SE  comer  of  section  23,  T 3 N,  R 17  W.  Locality  is  in  the 
Santa  Susana  quadrangle,  but  the  section  corner  is  in  the  Oat 
Mountain  quadrangle.  Bed  is  the  same  one  exposed  at  CSUN 
locality  468a.  [Shallow-marine  (transgressive)  facies.] 

516.  At  elevation  of  1975  ft.  (602  m)  on  north  side  of 
stream  bank  of  Las  Llajas  Canyon,  6500  ft.  (1981  m)  N67°W 
of  SE  comer  of  section  23,  T 3 N,  R 1 7 W.  Locality  is  in  the 
Santa  Susana  quadrangle,  but  the  section  corner  is  in  the  Oat 
Mountain  quadrangle.  Bed  is  the  same  one  exposed  at  CSUN 
locality  468b.  [Shallow-marine  (transgressive)  facies.] 

517.  At  elevation  of  1950  ft.  in  a small  canyon  bottom, 
between  the  letters  “n”  and  “y”  in  the  word  “Canyon,”  5100 
ft.  (1555  m)  N47°E  of  NE  comer  of  section  29,  T 3 N,  R 17 
W.  [Shallow-marine  (transgressive)  facies.] 

518.  At  elevation  of  2125  ft.  on  a small  ridge,  5475  ft. 
(1669  m)  N40°E  of  NE  comer  of  section  29,  T 3 N,  R 17  W. 
[Shallow-marine  (transgressive)  facies.] 

520.  At  elevation  of  2070  ft.,  6500  ft.  (1981  m)  N78°E  of 
NE  comer  of  section  29,  T 3 N,  R 17  W.  [Shallow-marine 
(transgressive)  facies.] 


521.  At  elevation  of  2 1 1 5 ft.  on  a ridge,  6360  ft.  ( 1 938  m) 
N88°E  of  NE  comer  of  section  29,  T 3 N,  R 1 7 W.  [Shallow- 
marine  (transgressive)  facies.] 

522.  Bed  131  ft.  (40  m)  stratigraphically  above  CSUN 
locality  37 1 , on  a traverse  bearing  S 1 4°W  from  locality  371. 
[Shallow-marine  (transgressive)  facies.] 

523.  At  elevation  of  2100  ft.  on  west  side  of  a canyon 
branching  south  of  Las  Llajas  Canyon,  5275  ft.  (1608  m) 
N70°W  of  SE  comer  of  section  23,  T 3 N,  R 17  W.  Locality 
is  in  the  Santa  Susana  quadrangle,  but  the  section  comer  is 
in  the  Oat  Mountain  quadrangle.  [Shallow-marine  (trans- 
gressive) facies.] 

524.  At  elevation  of  2035  ft.  on  ridge  along  south  side  of 
Las  Llajas  Canyon,  6375  ft.  (1943  m)  N72°W  of  SE  corner 
of  section  23,  T 3 N,  R 1 7 W.  Locality  is  in  the  Santa  Susana 
quadrangle,  but  the  section  corner  is  in  the  Oat  Mountain 
quadrangle.  [Shallow-marine  (transgressive)  facies.] 

525.  At  elevation  of  2050  ft.  on  a hillside,  6325  ft.  (1928 
m)  N74°W  of  SE  comer  of  section  23,  T 3 N,  R 1 7 W.  Locality 
is  in  the  Santa  Susana  quadrangle,  but  the  section  comer  is 
in  the  Oat  Mountain  quadrangle.  [Shallow-marine  (trans- 
gressive) facies.] 

526.  At  elevation  of  2055  ft.  on  a ridge,  2290  ft.  (698  m) 
N67°W  of  SE  comer  of  section  26,  T 3 N,  R 17  W.  Locality 
is  in  the  Santa  Susana  quadrangle,  but  the  section  corner  is 
in  the  Oat  Mountain  quadrangle.  [Shallow-marine  (trans- 
gressive) facies.] 

527a.  At  elevation  of  2075  ft.  on  a ridge,  2325  ft.  (709  m) 
N65°W  of  SE  comer  of  section  26,  T 3 N,  R 17  W.  Locality 
is  in  the  Santa  Susana  quadrangle,  but  the  section  corner  is 
in  the  Oat  Mountain  quadrangle.  [Shallow-marine  (trans- 
gressive) facies.] 

527b.  Bed  1 3 ft.  (4  m)  stratigraphically  above  locality  527a, 
on  a traverse  bearing  due  north  of  locality  527a.  [Shallow- 
marine  (transgressive)  facies.] 

528.  At  elevation  of  2185  ft.  on  a ridge,  1400  ft.  (427  m) 
north  and  2250  ft.  (686  m)  west  of  SE  comer  of  section  26, 
T 3 N,  R 17  W.  Locality  is  in  the  Santa  Susana  quadrangle, 
but  the  section  corner  is  in  the  Oat  Mountain  quadrangle. 
[Shallow-marine  (transgressive)  facies.] 

529a.  At  elevation  of  1400  ft.  on  east  side  of  stream  bank 
of  Las  Llajas  Canyon,  1 300  ft.  (396  m)  north  and  350  ft.  (107 
m)  east  of  SE  comer  of  section  29,  T 3 N,  R 1 7 W.  [Shallow- 
marine  (transgressive)  facies.] 

529b.  Bed  26.5  ft.  (7  m)  stratigraphically  above  locality 
529a,  on  a traverse  bearing  N27°E  of  locality  529a.  [Shallow- 
marine  (transgressive)  facies.] 

537.  At  elevation  of  1 500  ft.  on  west  side  of  an  amphi- 
theater on  west  side  of  Chivo  Canyon,  790  ft.  (241  m)  north 
and  2250  ft.  (686  m)  west  of  SE  comer  of  section  30,  T 3 N, 
R 17  W.  Locality  is  at  the  type  section  of  the  Llajas  Lor- 
mation.  [Shallow-marine  (transgressive)  facies.] 

538.  At  elevation  of  1 500  ft.  on  a ridge  on  west  side  of  Las 
Llajas  Canyon,  1500  ft.  (457  m)  north  and  350  ft.  (107  m) 
west  of  SE  comer  of  section  29,  T 3 N,  R 17  W.  [Shallow- 
marine  (transgressive)  facies.] 

539.  Bed  6 ft.  (1.8  m)  stratigraphically  above  locality  538, 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  63 


on  a traverse  bearing  N5°E  of  locality  538.  [Shallow-marine 
(transgressive)  facies.] 

540.  Bed  38  ft.  (11.5  m)  stratigraphically  below  CSUN 
locality  486,  on  a traverse  bearing  S28°E  from  locality  486. 
[Shallow-marine  (transgressive)  facies.] 

541.  At  elevation  of  1 325  ft.  on  north  bank  of  stream  bank 
of  Chivo  Canyon,  2800  ft.  (853  m)  north  and  1 175  ft.  (358 
m)  east  of  SE  corner  of  section  29,  T 3 N,  R 1 7 W.  [Outer 
shelf  and  slope  channel  facies.] 

542.  At  elevation  of  1315  ft.  on  ridge  near  mouth  of  Las 
Llajas  Canyon,  2200  ft.  (671  m)  north  and  800  ft.  (244  m) 
east  of  SW  comer  of  section  32,  T 3 N,  R 1 7 W.  [Interfingering 
coastal  alluvial-fan  facies  and  shallow-marine  (transgressive) 
facies.] 

543.  At  elevation  of  1460  ft.  on  divide  between  Bus  and 
Montgomery  Canyons,  600  ft.  (183  m)  south  and  1500  ft. 
(457  m)  east  of  NW  corner  of  section  28,  T 2 N,  R 18  W of 
the  USGS  7.5-minute  topographic  quadrangle  of  Thousand 
Oaks,  California  (1950),  photorevised  1967.  [Shallow-ma- 
rine (transgressive)  facies.] 

544.  At  elevation  of  1 230  ft.  on  a small  ridge,  1 025  ft.  (3 1 2 
m)  north  and  415  ft.  (127  m)  west  of  SE  corner  of  section 
19,  T 2 N,  R 18  W of  the  USGS  7.5-minute  topographic 
quadrangle  of  Thousand  Oaks,  California  (1950),  photore- 
vised 1967.  [Shallow-marine  (regressive)  facies.] 

545.  At  elevation  of  1 120  ft.  on  east  side  of  Bus  Canyon, 
580  ft.  (1 77  m)  south  and  14 10  ft.  (430  m)  west  of  NE  comer 
of  section  28,  T 2 N,  R 18  W of  the  USGS  7.5-minute 
topographic  quadrangle  of  Thousand  Oaks,  California  (1950), 
photorevised  1967.  [Interfingering  coastal  alluvial-fan  facies 
and  shallow-marine  (transgressive)  facies.] 

546.  At  elevation  of  1 500  ft.  on  east  side  of  divide  between 
Bus  and  Trough  Canyons,  700  ft.  (213  m)  south  and  200  ft. 
(61  m)  east  of  NW  corner  of  section  27,  T 2 N,  R 18  W of 
the  USGS  7.5-minute  topographic  quadrangle  of  Thousand 
Oaks,  California  (1950),  photorevised  1967.  [Shallow-ma- 
rine (transgressive)  facies.] 

547.  At  elevation  of  1 0 1 0 ft.  on  east  side  of  a roadcut,  650 
ft.  (198  m)  south  of  NE  corner  of  section  21,  T 2 N,  R 18 
W of  the  USGS  7.5-minute  topographic  quadrangle  of  Thou- 
sand Oaks,  California  (1950),  photorevised  1967.  [Shallow- 
marine  (regressive)  facies.] 

548.  At  elevation  of  947  ft.  on  south  side  of  Simi  Arroyo 
just  above  the  streambed,  1 100  ft.  (335  m)  north  and  1750 
ft.  (553  m)  west  of  SE  corner  of  section  12,  T 2 N,  R 18  W. 
Locality  is  equivalent  to  UCLA  locality  5837.  [Shallow-ma- 
rine (transgressive)  facies.] 

700.  At  elevation  of  1 100  ft.  along  east  side  of  road  in  Bus 
Canyon,  750  ft.  (229  m)  south  and  1500  ft.  (457  m)  west  of 
NE  comer  of  section  28,  T 2 N,  R 18  W of  the  USGS  7.5- 
minute  topographic  quadrangle  of  Thousand  Oaks,  Califor- 
nia (1950),  photorevised  1967.  [Interfingering  coastal  allu- 
vial-fan facies  and  shallow-marine  (transgressive)  facies.] 

701.  At  elevation  of  1 160  ft.  along  east  side  of  Bus  Canyon, 
800  ft.  (244  m)  south  and  1400  ft.  (427  m)  west  of  NE  corner 
of  section  28,  T 2 N,  R 18  W of  the  USGS  7.5-minute 
topographic  quadrangle  of  Thousand  Oaks,  California  (1950), 
photorevised  1967.  [Interfingering  coastal  alluvial-fan  facies 
and  shallow-marine  (transgressive)  facies.] 


702.  At  elevation  of  1420  ft.  along  a ridge,  900  ft.  (274  m) 
south  and  680  ft.  (207  m)  west  of  NE  corner  of  section  28, 
T 2 N,  R 18  W of  the  USGS  7.5-minute  topographic  quad- 
rangle of  Thousand  Oaks,  California  (1950),  photorevised 
1967.  Locality  is  equivalent  to  UCMP  locality  7193  and 
UCLA  locality  7070.  [Interfingering  coastal  alluvial-fan  fa- 
cies and  shallow-marine  (transgressive)  facies.] 

703.  At  elevation  of  1430  ft.  along  a ridge,  780  ft.  (238  m) 
south  and  580  ft.  (177  m)  west  of  NE  corner  of  section  28, 
T 2 N,  R 18  W of  the  USGS  7.5-minute  topographic  quad- 
rangle of  Thousand  Oaks,  California  (1950),  photorevised 
1 967.  Locality  is  a few  meters  stratigraphically  above  locality 
702.  [Lowermost  part  of  shallow-marine  (transgressive)  fa- 
cies.] 

704.  At  elevation  of  1 320  ft.  along  divide  between  Bus  and 
Trough  Canyons,  180  ft.  (55  m)  south  and  90  ft.  (27  m)  west 
of  NE  comer  of  section  28  of  the  USGS  7.5-minute  topo- 
graphic quadrangle  of  Thousand  Oaks,  California  (1950), 
photorevised  1967.  [Shallow-marine  (transgressive)  facies.] 

705.  At  elevation  of  1240  ft.  along  a dip-slope  just  south 
of  the  mouth  of  Las  Llajas  Canyon,  2450  ft.  (747  m)  north 
and  450  ft.  (137  m)  east  of  SW  corner  of  section  32,  T 3 N, 
R 17  W.  [Interfingering  coastal  alluvial-fan  facies  and  shal- 
low-marine (transgressive)  facies.] 

706.  At  elevation  of  1250  ft.  along  a small  ridge  on  east 
side  of  Chivo  Canyon  where  it  bends  sharply  to  the  northeast, 
700  ft.  (213  m)  north  and  590  ft.  (180  m)  west  of  SE  corner 
of  section  30,  T 3 N,  R 1 7 W.  [Interfingering  coastal  alluvial- 
fan  facies  and  shallow-marine  (transgressive)  facies.] 

CAS  LOCALITIES 

25.  On  east  bank  of  Little  River  at  its  confluence  with 
Umpqua  River,  near  center  of  section  19,  T 26  S,  R 3 W. 
Roseburg  quadrangle,  Douglas  County,  Oregon. 

364.  Exact  location  unknown.  Aliso  Creek  (now  known  as 
Devil  Canyon),  section  25,  T 3 N,  R 17  W,  near  Chatsworth, 
California,  Los  Angeles  County. 

372.  Exact  location  unknown.  Aliso  Canyon  (now  known 
as  Devil  Canyon),  section  23,  T 3 N,  R 17  W,  near  Chats- 
worth, California,  Los  Angeles  County. 

393.  Devil  Canyon,  SE  'A  of  the  NW  'A  of  section  26,  T 3 
N,  R 17  W,  Santa  Susana  quadrangle,  California. 

711.  East  side  of  Grapevine  Creek  near  point  where  it 
enters  valley  floor,  Tejon  quadrangle,  Kern  County,  Califor- 
nia. 

792.  West  side  of  Tecuya  Creek,  about  1 mile  south  of 
where  stream  flows  out  on  valley  floor,  Tejon  quadrangle, 
Kem  County,  California. 

HERMAN  SCHYMICZEK  LOCALITY 

HS-A-1 1(4).  Bed  92  ft.  (28  m)  stratigraphically  above  CSUN 
locality  4691,  on  a traverse  bearing  due  north  from  locality 
4691. 

LACMIP  LOCALITY 

461-B.  On  the  northern  slope  of  a small  canyon  intersecting 
Las  Llajas  Canyon  from  the  east.  It  is  200  ft.  (61  m)  from 


64  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


the  top  of  the  Llajas  Formation,  Las  Llajas  Canyon,  Santa 
Susana  Mountains,  Ventura  County,  California. 

SU  LOCALITY 

2696.  Chivo  Canyon,  5 km  N20°E  of  Bench  Mark  961  at 
Santa  Susana,  Santa  Susana  quadrangle,  Ventura  County, 
California. 

UCMP  LOCALITIES 

337.  About  5 km  south  of  Martinez,  on  the  east  side  of 
the  road  to  Walnut  Creek,  Concord  quadrangle.  Contra  Costa 
County,  California. 

452.  On  road  in  Grapevine  Creek,  SW  ‘A  of  the  SE  ‘A  of 
section  20,  T 10  N,  R 19  W,  Tejon  quadrangle,  Kern  County, 
California. 

672.  South  portion  of  crest  of  Parson’s  Peak,  SE  ‘A  of  the 
NW  'A  of  section  24,  T 18  S,  R 14  E,  Coalinga  quadrangle, 
Fresno  County,  California. 

1427.  SW  ‘A  of  section  1 1,  NE  of  Wall  Point,  south  side 
of  Mount  Diablo,  California. 

1817.  Opposite  the  place  where  Urruttia  Canyon  enters 
Salt  Creek,  100  ft.  (30  m)  up  fourth  small  draw  from  west 
end  of  ridge,  SW  ‘A  of  the  NW  ‘A  of  section  15,  T 18  S,  R 
14  E,  Coalinga  quadrangle,  Fresno  County,  California. 

1853.  Marysville  Buttes,  N Vi  of  section  28,  T 16  N,  R 1 
E,  Marysville  Buttes  quadrangle,  Sutter  County,  California. 

2226.  Longitude  1 17°14'W,  latitude  33°50'N,  SE  of  Sole- 
dad  Mountain,  north  of  Ladrillo  Station,  Southern  Pacific 
Railroad,  Rose  Canyon,  La  Jolla  quadrangle,  San  Diego 
County,  California. 

2287.  West  side  of  Domengine  Canyon,  SW  comer  of  the 
SW  'A  of  the  SE  ‘A  of  section  29,  T 18  S,  R 15  E,  Fresno 
County,  California. 

2295.  Longitude  120. 1°W,  latitude  36.1°N,  near  Domen- 
gine Ranch,  Fresno  County,  California. 

3296.  Exact  location  unknown.  Aliso  Canyon  (now  known 
as  Devil  Canyon),  SE  ‘A  of  the  SE  ‘A  of  section  23,  T 3 N,  R 
1 7 W,  Santa  Susana  quadrangle,  Ventura  County,  California. 

3304.  Exact  location  unknown.  Aliso  Canyon  (now  known 
as  Devil  Canyon,  section  36,  T 3 N,  R 17  W,  Santa  Susana 
quadrangle,  Los  Angeles  County,  California. 

3310.  Exact  location  unknown.  Probably  Simi  Hills,  Santa 
Susana  quadrangle,  California. 

3976.  Seven-eighths  of  a mile  (1.4  km)  north  of  Ladrillo 
Station,  Southern  Pacific  Railroad,  in  first  canyon  east  of 
Rose  Canyon,  elevation  259  ft..  La  Jolla  quadrangle,  San 
Diego  County,  California. 

3981.  At  15  m above  high-tide  level  in  small  gully  0.4  km 
south  of  mouth  of  Soledad  Valley,  La  Jolla  quadrangle,  San 
Diego  County,  California. 

3989.  0.2  mi.  (0.3  km)  north  of  the  top  of  the  “y”  of 
“Tecolote  Valley,”  on  the  east  side  of  the  valley,  elevation 
100  ft.,  La  Jolla  quadrangle,  San  Diego  County,  California. 

3990.  On  the  east  side  of  canyon  in  bottom  of  Rose  Creek, 
0.3  mi.  (0.5  km)  east  of  “t”  of  “Soledad  Mountain,”  La  Jolla 
quadrangle,  San  Diego  County,  California. 

3993.  In  bottom  of  Rose  Creek  where  creek  makes  a strong 
bend  to  west,  0.2  mi.  (0.3  km)  south  of  Bench  Mark  176,  2 


mi.  (3.2  km)  east  of  La  Jolla,  La  Jolla  quadrangle,  San  Diego 
County,  California. 

4169.  About  500  ft.  (152  m)  east  of  ranch  house  in  Big  Tar 
Canyon  on  east  line  of  section  18,  near  point  where  road 
crosses  creek,  Garza  Peak  quadrangle.  Kings  County,  Cali- 
fornia. 

4170.  On  west  side  of  Big  Tar  Canyon,  where  it  crosses 
the  Eocene  section,  Garza  Peak  quadrangle,  Kings  County, 
California. 

4175.  No  locality  data  available.  Probably  Domengine 
Ranch  area,  north  of  Coalinga,  Domengine  Ranch  quadran- 
gle, Fresno  County,  California. 

5062.  In  sea  cliff  south  of  mouth  of  Soledad  Valley,  due 
west  of  midpoint  between  “P”  and  “u”  of  Pueblo,”  La  Jolla 
quadrangle,  San  Diego  County,  California. 

5069.  0.4  inches  SSE  of  the  “a”  in  “Morena”  in  a branch 
canyon  of  Tecolote  Valley  to  the  north,  elevation  1 50  ft.,  La 
Jolla  quadrangle,  San  Diego  County,  California. 

5080.  North  side  of  Soledad  Valley,  at  McGonigle  Canyon, 
NE  ‘A  of  section  36,  T 14  S,  R 4 W,  La  Jolla  quadrangle,  San 
Diego  County,  California. 

5085.  2.62  inches  north  of  the  top  of  the  “S”  of  “Soledad 
Mountain,”  on  the  north  side  of  the  creek,  on  a small  ridge 
formed  by  the  creek  and  sea  cliff,  elevation  75  ft..  La  Jolla 
quadrangle,  San  Diego  County,  California. 

5089.  300  ft.  (91  m)  north  of  the  Scripps  Institution  pier, 
in  the  conglomerate  above  the  mudstone.  In  the  sea  cliff, 
elevation  10  ft.,  La  Jolla  quadrangle,  San  Diego  County, 
California. 

7000.  Exact  location  unknown.  Las  Llajas  Canyon,  in  first 
canyon  on  north  side  of  road,  Santa  Susana  quadrangle,  Cal- 
ifornia. 

7002.  At  elevation  of  1750  ft.,  in  road  near  top  of  last 
western  spur  which  extends  northward  out  of  Las  Llajas  Can- 
yon. In  shales  striking  north  just  south  of  east-west  fault 
which  is  made  prominent  by  the  Meganos  Conglomerate 
which  parallels  it  on  the  north  side,  Santa  Susana  quadrangle, 
Ventura  County. 

7003.  At  elevation  of  1 760  ft.  in  sandstone  exposed  in  deep 
washout  on  south  side  of  second  large  canyon  which  enters 
Llajas  Canyon  from  the  east.  Washout  is  about  100  yards 
east  of  mouth  of  canyon.  Fossils  are  found  in  ledge  on  west 
side  of  washout  and  near  top.  Sandstone  here  dips  with  slope 
of  canyon  side  and  from  west  wall  of  east-west  fault  which 
crosses  Llajas  Canyon  south  of  this  locality,  Santa  Susana 
quadrangle,  Ventura  County. 

7004.  About  300  ft.  (91  m)  east  of  locality  7003  in  next 
small  canyon  which  enters  Llajas  Canyon  from  the  east  just 
south  of  the  most  northerly  extension  of  the  1 500-ft.  contour, 
Santa  Susana  quadrangle,  Ventura  County.  Locality  is  equiv- 
alent to  CSUN  locality  374. 

7005.  No  locality  data  available.  Probably  Las  Llajas  Can- 
yon area,  Santa  Susana  quadrangle,  California. 

7015.  About  5500  ft.  (1676  m)  S69°E  of  Bench  Mark  at 
Santa  Susana,  Santa  Susana  quadrangle,  Ventura  County. 

7182.  Boulder  from  second  draw  on  east  side  of  Live  Oak 
Canyon,  Tejon  Ranch,  Kern  County,  California. 

7193.  About  600  ft.  (183  m)  east  of  and  apparently  slightly 
stratigraphically  beneath  locality  7194,  which  is  along  the 


Contributions  in  Science,  Number  350 


Squires:  Sinii  Valley  Eocene  Mollusks  65 


top  of  a 1400-ft.  ridge  extending  NW  from  third  main  ridge 
west  of  Runkle’s  Ranch  on  the  ridge  on  which  there  is  an 
abandoned  well  and  road  leading  up  to  it,  Simi  Valley,  Cal- 
ifornia. Locality  is  equivalent  to  CSUN  locality  702  and 
UCLA  locality  7070. 

7195.  In  the  creek  bed  about  60  ft.  (18  m)  north  of  the 
second  falls  or  300  ft.  (91  m)  north  of  the  mouth  of  the  first 
small  draw  which  enters  Las  Llajas  Canyon  west  of  the  point 
where  the  Meganos  Conglomerate  crosses  the  road,  Santa 
Susana  quadrangle,  Ventura  County,  California. 

7200.  No  locality  data  available.  Probably  Live  Oak  Can- 
yon area,  Tejon  Ranch,  Kern  County,  California. 

A-661.  On  east  bank  of  Little  River  between  the  highway 
bridge  and  the  first  bend  of  the  stream  east  of  the  junction 
with  the  North  Umpqua  River,  center  of  section  19,  T 26  S, 
R 3 W,  Douglas  County,  Oregon. 

A-819.  Lowest  reef  bed  on  side  of  hill  just  east  of  and 
above  first  saddle  south  of  Big  Tar  Canyon,  Garza  Peak 
quadrangle.  Kings  County,  California. 

A-993.  Second  draw  past  Marrland  Canyon  (now  known 
as  Las  Llajas  Canyon)  at  second  small  falls  up  draw  approx- 
imately 600  ft.  (183  m),  Santa  Susana  quadrangle,  Ventura 
County,  California. 

A-994.  About  225  to  300  ft.  (69  to  9 1 m)  down  the  canyon 
from  locality  A-993  on  west  side  of  canyon  about  60  ft.  (18 
m)  from  streambed,  Santa  Susana  quadrangle,  Ventura 
County,  California. 

A-1003.  Exact  location  unknown.  Pine  Canyon,  Mount 
Diablo,  Contra  Costa  County,  California. 

A-1027.  Valdes  Ranch,  on  branch  of  Silver  Creek,  Valle- 
citos,  center  of  east  part  of  SW  *A  of  section  4,  T 16  S,  R 12 
E.  Approximately  where  1 20°40'  parallel  crosses  most  north- 
erly intermittent  stream  indicated  on  section  4,  Panoche 
quadrangle,  Fresno  County,  California. 

A-1219.  Base  of  Domengine  Formation  on  west  side,  near 
top,  of  long  ridge  extending  NW  of  2126-ft.  hill  on  line  be- 
tween sections  9 and  16,  T 19  S,  R 1 5 E,  Domengine  Ranch 
quadrangle,  California. 

A-1280.  Near  center  of  north  edge  of  section  20,  on  hill 
immediately  south  of  point  where  the  Big  Tar-McLure  Val- 
ley road  crosses  saddle  at  head  of  steam  running  into  McLure 
Valley,  45  ft.  (14  m)  below  uppermost  fossiliferous  layer, 
Garza  Peak  quadrangle.  Kings  County,  California. 

A-3042.  On  ridge  in  blue  clay  shale  member  above  con- 
glomerate member  at  base  of  Llajas  Formation,  center  of  SE 
■A  of  the  SE  'A  of  section  10,  T 3 N,  R 17  W,  Santa  Susana 
quadrangle,  Los  Angeles  County,  California. 

UCLA  LOCALITY 

6616.  West  end  of  ridge  south  of  Las  Llajas  Canyon,  2525 
ft.  (770  m)  north  and  575  ft.  (175  m)  east  of  SW  corner  of 
section  32,  T 3 N,  R 1 7 W,  Santa  Susana  quadrangle,  Ventura 
County,  California. 

UCR  LOCALITY 

4865.  In  a roadcut  on  the  southwest  side  of  California  State 
Highway  78,  5.2  km  SE  of  Vista  and  4.8  km  NW  of  San 


Marcos,  at  longitude  117°12'39"W,  latitude  33°09'47"N,  in 
northern  San  Diego  County,  California. 

UO  LOCALITY 

144.  About  2.5  mi.  (4  km)  up  Fall  Creek  from  Little  River 
on  Mathews  farm  in  NE  'A  of  section  6,  T 27  S,  R 3 W, 
Douglas  County,  Oregon. 

USGS  LOCALITIES 

4617.  On  SW  flank  of  Reef  Ridge,  north  of  McLure  Valley, 
2‘A  mi.  (3.6  km)  SSE  of  El  Cerrito  oil  well,  in  section  27,  T 
23  S,  R 1 7 E,  Cholame  quadrangle.  Kings  County,  California. 

4619.  North  of  Coalinga,  15  mi.  (24  km)  SW  of  Domen- 
gine’s  Ranch,  T 18  S,  R 15  E,  Coalinga  quadrangle,  Fresno 
County,  California. 

UW  LOCALITIES 

329.  On  north  bank  of  the  Cowlitz  River  at  bend  1.5  to 
2.5  km  east  ofVader,  section  28,  T 1 N,  R 2 W,  Lewis  County, 
Washington. 

358.  Joice  Station,  ‘A  mi.  (0.4  km)  east  of  Tongue  Point 
Railroad,  Port  Crescent,  section  22,  T 3 1 N,  R 8 W,  Clallam 
County,  Washington. 

ACKNOWLEDGMENTS 

Acknowledgment  is  made  to  the  donors  of  The  Petroleum 
Research  Fund,  administered  by  the  American  Chemical  So- 
ciety, for  support  of  this  research  under  grant  number  PRF 
1 1472-B2. 

The  following  ranchers  kindly  permitted  access  to  the  field: 

G.  Haigh,  W.  Haigh,  D.  Poe,  G.  Boyle,  and  J.  Runkle.  Sincere 
thanks  are  extended  to  the  following  California  State  Uni- 
versity, Northridge,  geology  students  for  their  assistance  in 
the  field:  D.  Advocate,  D.  Carpenter,  G.  Davis,  S.  Drewry, 
T.  Santochi,  M.  Stettner,  and  G.  Wood.  G.  Davis  helped  in 
photographing  the  fossils. 

The  following  people  kindly  provided  the  loan  of  speci- 
mens: F.  J.  Collier,  United  States  National  Museum  of  Nat- 
ural History;  F.  A.  Jenkins,  Loyola  Marymount  University; 

H.  A.  Lowenstam,  California  Institute  of  Technology;  E.  Nes- 
bitt and  J.  Peck,  University  of  California,  Berkeley;  P.  U. 
Rodda,  California  Academy  of  Sciences;  and  L.  R.  Saul, 
University  of  California,  Los  Angeles. 

I am  grateful  to  A.  G.  Beu  (New  Zealand  Geological  So- 
ciety), C.  R.  Givens  (Nicholls  State  University),  and  L.  R. 
Saul  for  their  valuable  comments  on  molluscan  taxonomy 
and  identification.  L.  R.  Saul  also  helped  greatly  in  the  doc- 
umentation of  the  early  Eocene  molluscan  fauna  of  the  Llajas 
Formation.  B.  J.  Welton  (Chevron  Research,  California) 
kindly  identified  the  shark  teeth.  M.  Filewicz  and  M.  Hill, 
III  (Union  Oil  of  California)  shared  their  calcareous  nan- 
nofossil  data.  D.  R.  Lindberg  (University  of  California, 
Berkeley)  and  T.  Susuki  (University  of  California,  Los  An- 
geles) provided  information  as  to  the  location  of  some  fossil 
localities.  R.  Simpson  and  E.  C.  Wilson  (Los  Angeles  County 
Museum)  gave  most  helpful  editorial  comments.  C.  R.  Giv- 


66  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


ens  and  W.  J.  Zinsmeister  (Purdue  University)  critically  re- 
viewed the  manuscript,  and  their  suggestions  greatly  im- 
proved it. 

LITERATURE  CITED 

Abbott,  R.T.  1968a.  Seashells  of  North  America.  Golden 
Press,  New  York.  280  pp. 

. 1968b.  The  helmet  shells  of  the  world  (Cassidae), 

Part  1.  Indo-Pacific  Mollusca  2(9):  15-200. 

. 1974.  American  seashells,  2nd  ed.  Van  Nostrand 

Reinhold  Company,  New  York.  663  pp. 

Abbott,  R.T.,  and  S.P.  Dance.  1982.  Compendium  of  sea 
shells.  E.P.  Dutton,  New  York.  41 1 pp. 

Adams,  Arthur.  1850.  In  Sowerby,  G.B.,  Thesaurus  con- 
chyliorum,  or  monographs  of  genera  of  shells,  vol.  2. 
London. 

. 1 864.  On  the  species  of  Neara  found  in  the  seas  of 

Japan.  Annales  Magasin  Naturelle  Histoire,  ser.  3,  13: 
206-209. 

Adams,  Henry,  and  Arthur  Adams.  1853-1858.  The  genera 
of  recent  Mollusca;  arranged  according  to  their  organi- 
zation. 3 vols.  John  Van  Voorst,  London.  661  pp. 

Addicott,  W.O.  1970.  Latitudinal  gradients  in  Tertiary 
molluscan  faunas  of  the  Pacific  Coast.  Palaeogeography, 
Palaeoclimatology,  Palaeoecology  8:287-312. 

Advocate,  D.M.  1982.  Depositional  environments  of  the 
Maniobra  Formation,  northwestern  Orocopia  Moun- 
tains, Riverside  County,  California.  California  State 
University,  Northridge,  Unpublished  Master  of  Science 
thesis.  130  pp. 

Agassiz,  L.J.R.  1836.  Memoires  de  la  Societe  Science  Na- 
turelle Neuchatel,  vol.  1.  Paris. 

. 1838-1843.  Recherches  sur  les  poissons  fossiles. 

Memoires  de  la  Societe  Science  Naturelle  Neuchatel  3: 
73-357. 

Aldrich,  T.H.  1886.  Preliminary  report  on  the  Tertiary 
fossils  of  Alabama  and  Mississippi.  Geological  Survey 
of  Alabama,  Bulletin  1 (1):  15-60,  6 pis. 

. 1911.  New  Eocene  fossils  from  the  southern  Gulf 

States.  Bulletins  of  American  Paleontology  5(22).  24  pp. 

Anderson,  F.M.  1905.  A stratigraphic  study  in  the  Mount 
Diablo  Range  of  California.  California  Academy  of  Sci- 
ences, Proceedings,  ser.  3,  2(2):  1 55-248. 

Anderson,  F.M. , and  G.D.  Hanna.  1925.  Fauna  and  strati- 
graphic relations  of  the  Tejon  Eocene  at  the  type  locality 
in  Kern  County,  California.  California  Academy  of  Sci- 
ences, Occasional  Papers  1 1:1-249. 

. 1928.  Solen  novacularis,  a name  for  an  Eocene 

fossil  from  California.  The  Nautilus  42(2):65-66. 

Anton,  H.E.  1839.  Verzeichnissder Conchylien  welche  sich 
in  der  Sammlung  von  Herman  Eduard  Anton  befinden. 
Halle.  110  pp. 

Armentrout,  J.M.  1975.  Molluscan  biostratigraphy  of  the 
Lincoln  Creek  Formation,  southwest  Washington.  Pages 
14-28  in  Weaver,  D.E.,  G.R.  Homaday,  and  Ann  Tip- 
ton,  eds.,  Future  energy  horizons  of  the  Pacific  Coast; 
Paleogene  symposium  and  selected  technical  papers. 


American  Association  of  Petroleum  Geologists,  Society 
of  Economic  Paleontologists  and  Mineralogists,  and  So- 
ciety of  Economic  Geologists,  Pacific  Sections,  Annual 
Meeting. 

Arnold,  Ralph.  1907.  New  and  characteristic  species  of 
fossil  mollusks  from  the  oil-bearing  Tertiary  formations 
of  southern  California.  United  States  National  Museum, 
Proceedings  32:525-546. 

. 1909.  Environment  of  the  Tertiary  faunas  of  the 

Pacific  Coast  of  the  United  States.  Journal  of  Geology 
17:509-533. 

. 1910.  Paleontology  of  the  Coalinga  district,  Fresno 

and  Kings  Counties,  California.  United  States  Geolog- 
ical Survey,  Bulletin  396:1-173. 

Arnold,  Ralph,  and  H.  Hannibal.  1913.  The  marine  Ter- 
tiary stratigraphy  of  the  North  Pacific  coast  of  America. 
American  Philosophical  Society,  Proceedings  52:559- 
605. 

Baldwin,  E.M.  1974.  Eocene  stratigraphy  of  southwestern 
Oregon.  Oregon  Department  of  Geology  and  Mineral 
Industries,  Bulletin  83.  40  pp. 

Basterot,  B.  de.  1825.  Description  geologique  du  bassin 
tertiaire  du  sud-ouest  de  la  France,  premiere  partie  com- 
prenant  les  observations  generates  sur  les  Mollusques 
fossiles.  Paris  Societe  Histoire  Naturelle,  Memoire  2:1- 
100. 

Bell,  Thomas.  1858.  A monograph  of  the  fossil  malacos- 
tracous  of  Great  Britain.  Part  1 , Crustacea  of  the  London 
Clay.  Palaeontographical  Society  of  London.  44  pp. 

Bellardi,  Luigi.  1872-1890.  I molluschi  dei  terreni  terziari 
del  piemonte  e della  Liguria.  Musei  di  Zoologia  ed  An- 
atomia  comparata  della  R.  Universita  di  Torino,  Bol- 
lettino,  pts.  1-5. 

Bentson,  Herdis.  1940.  A systematic  study  of  the  fossil 
gastropod  Exilia.  University  of  California,  Publications 
in  Geological  Sciences  25(5):  1 99-238. 

Berggren,  W.A.,  D.V.  Kent,  and  J.J.  Flynn.  Paleogene  geo- 
chronology and  chronostratigraphy.  In  Snelling,  N.J., 
ed.,  Geochronology  and  the  geological  record.  Geolog- 
ical Society  of  London,  Special  Paper  (in  press). 

Berthiaume,  S.A.  1938.  Orbitoids  from  the  Crescent  For- 
mation (Eocene)  of  Washington.  Journal  of  Paleontology 
12:494-497. 

Blainville,  H.M.D.  de.  1814.  Memoire  sur  la  classification 
methodique  des  animaux  mollusques,  et  etablissement 
d’une  nouville  consideration  pour  y parvenir.  Societe 
Philomathematiques  Paris,  Bulletin,  pp.  175-180. 

. 1816-1830.  Vers  et  zoophytes.  In  Dictionnaire  des 

sciences  naturelles.  Part  2,  Regne  organise.  60  vols. 

Blondeau,  Alphonse,  and  E.E.  Brabb.  1983.  Large  fora- 
minifers  of  Eocene  age  from  the  Coast  Ranges  of  Cali- 
fornia. Pages  41-48  in  Brabb,  E.E.,  ed..  Studies  in  Ter- 
tiary stratigraphy  of  the  California  Coast  Ranges.  United 
States  Geological  Survey,  Professional  Paper  1213. 

Bom,  Ignatius.  1780.  Testacea  Musei  Caesarei  Vindobo- 
nensis.  Joannis  Pauli  Kraus,  Vienna.  442  pp. 

Bronn,  H.G.  1837-1838.  Lethaea  geognostica  Oder  Abbil- 
dungen  und  Beschreibungen  der  fur  die  Gebirgsforma- 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  67 


tionen  bezeichnenden  Versteinerungen.  Schweizerbart, 
Stuttgart.  1346  pp. 

. 1849.  Handbuch  der  Geschichte  der  Natur.  Index 

Palaeontologicus,  vol.  3 (pt.  3).  Stuttgart.  976  pp. 

Bruguiere,  J.G.  1789-1816.  Encyclopedic  methodique  ou 
par  ordre  et  matieres.  Histoire  naturelle  des  Vers,  des 
Mollusques.  758  pp. 

Chavan,  Andre.  1969.  Superfamily  Lucinacea  Fleming, 
1 828.  Part  N,  Mollusca  6,  Bivalvia,  vol.  2,  pp.  491-518 
in  Moore,  R.C.,  ed..  Treatise  on  invertebrate  paleon- 
tology. University  of  Kansas  Press,  Lawrence. 

Chemnitz,  J.H.  1780-1795.  Neues  systematisches  conchy- 
lien  cabinet,  vols.  4-1 1.  Bauer  and  Raspe,  Nurnberg. 

Children,  J.G.  1822-1824.  Lamarck’s  genera  of  shells, 
translated  from  the  French.  Quarterly  Journal  of  Science, 
vols.  14-16. 

Clark,  B.L.  1918.  The  San  Lorenzo  Series  of  middle  Cal- 
ifornia. University  of  California,  Publications  in  Geo- 
logical Sciences  1 1(2):45-234. 

. 1921.  The  stratigraphic  and  faunal  relationships  of 

the  Meganos  Group,  middle  Eocene  of  California.  Jour- 
nal of  Geology  29: 1 25-165. 

. 1926.  The  Domengine  horizon,  middle  Eocene  of 

California.  University  of  California,  Publications  in 
Geological  Sciences  1 6(5):99—  1 18. 

. 1929.  Stratigraphy  and  faunal  horizons  of  the  Coast 

Range  of  California.  Privately  published.  132  pp. 

. 1934.  A new  genus  and  two  new  species  of  La- 

mellibranchiata  from  the  middle  Eocene  of  California. 
Journal  of  Paleontology  8(3):270— 272,  pi.  37. 

. 1938.  Fauna  from  the  Markley  Formation  (upper 

Eocene)  on  Pleasant  Creek,  California.  Geological  So- 
ciety of  America,  Bulletin  49(5):683— 730. 

. 1942.  New  middle  Eocene  gastropods  from  Cali- 
fornia. Journal  of  Paleontology  1 6(  1 ):  1 16-1 19. 

Clark,  B.L.,  and  D.K.  Palmer.  1923.  Revision  of  the  Ri- 
mella- like  gastropods  from  the  West  Coast  of  North 
America.  University  of  California,  Publications  in  Geo- 
logical Sciences  14(7):277-288. 

Clark,  B.L.,  and  H.E.  Vokes.  1936.  Summary  of  marine 
Eocene  sequence  of  western  North  America.  Geological 
Society  of  America,  Bulletin  47:851-878. 

Clark,  B.L.,  and  Woodford,  A.O.  1927.  The  geology  and 
paleontology  of  the  type  section  of  the  Meganos  For- 
mation (lower  middle  Eocene)  of  California.  University 
of  California,  Publications  in  Geological  Sciences  17(2): 
63-142. 

Cole,  W.S.  1958.  Names  and  variations  in  certain  Amer- 
ican larger  Foraminifera,  particularly  the  Discocyclinids. 
Bulletins  of  American  Paleontology  38:41 1-429. 

. 1964.  Discocyclinidae.  Part  C,  Protista  2,  vol.  2, 

pp.  712-717  in  Moore,  R.C.,  ed..  Treatise  on  inverte- 
brate paleontology.  University  of  Kansas  Press,  Law- 
rence. 

Cole,  W.S.,  and  E.R.  Applin.  1964.  Problems  of  the  geo- 
graphic and  stratigraphic  distribution  of  American  mid- 
dle Eocene  larger  Foraminifera.  Bulletin  of  American 
Paleontology  47.  48  pp. 


Compagno,  L.J.U.  1973.  Interrelationships  of  living  elas- 
mobranchs.  Pages  1 5-6 1 in  Greenwood,  P.H.,  R.S.  Miles, 
and  C.  Patterson,  eds.,  Interrelationships  of  fishes.  Sup- 
plement Number  1,  Journal  of  the  Linnean  Society  of 
London,  Zoology.  Vol.  53. 

Conrad,  T.A.  1832.  Fossil  shells  of  the  Tertiary  formations 
of  North  America.  Vol.  1,  no.  2.  Privately  published.  17 

pp. 

. 1833.  On  some  new  fossil  and  Recent  shells  of  the 

United  States.  American  Journal  of  Science  and  Arts 
23(2):339-346. 

. 1834.  Observations  on  the  Tertiary  and  more  re- 
cent formations  of  a portion  of  the  southern  states.  Ap- 
pendix, descriptions  of  new  Tertiary  fossils  from  the 
southern  states.  Academy  of  Natural  Sciences  of  Phila- 
delphia, Journal  7: 1 16-157. 

. 1855.  Report  on  the  fossil  shells  collected  in  Cal- 
ifornia by  W.P.  Blake.  Pages  5-20  in  Preliminary  geo- 
logical report  of  W.P.  Blake,  United  States  33rd  Con- 
gress, 1st  session.  House  Executive  Document  129. 

. 1857.  Descriptions  of  the  fossil  shells.  Pages  317- 

329  in  Williamson,  R.S.,  Report  of  explorations  in  Cal- 
ifornia for  railroad  routes.  United  States  33rd  Congress, 
2nd  session.  Senate  Executive  Document  78  and  House 
Executive  Document  91,  vol.  5,  pt.  2,  appendix  2. 

. 1860.  Description  of  new  species  of  Cretaceous  and 

Eocene  fossils  of  Mississippi  and  Alabama.  Academy  of 
Natural  Sciences  of  Philadelphia,  Journal,  ser.  2,  4:279- 
296. 

. 1865.  Catalogue  of  the  Eocene  and  Oligocene  tes- 

tacea  of  the  United  States.  American  Journal  of  Con- 
chology  1:1-35. 

. 1866.  Check  list  of  the  invertebrate  fossils  of  North 

America.  Eocene  and  Oligocene.  Smithsonian  Miscel- 
laneous Collections  7(200):  1-41. 

. 1867.  Descriptions  of  new  genera  and  species  of 

Miocene  shells,  with  notes  on  other  fossil  and  recent 
species.  American  Journal  of  Conchology  3:257-270. 

Cooper,  J.G.  1894.  Catalogue  of  Californian  fossils,  parts 
2-5.  California  State  Mining  Bureau,  Bulletin  4:5-65. 

Cossmann,  A.E.M.  1888.  Note  rectificative  sur  la  nomen- 
clature d’un  genre  de  coquilles  fossiles.  Journal  Con- 
chyliologie  36:335. 

. 1889.  Catalogue  illustre  des  coquilles  fossiles  de 

l’Eocene  des  environs  de  Paris.  Gastropodes.  Annales 
de  la  Societe  Royale  Malacologique  de  Belgique,  vol.  24. 

. 1895-1925.  Essais  de  paleoconchologie  comparee. 

1 3 livraisons.  Les  Presses  Universitaires  de  France,  Paris. 

Costa,  E.M.  da.  1776.  Elements  of  conchology:  or  an  in- 
troduction to  the  knowledge  of  shells.  London.  318  pp. 

. 1778.  Historia  naturalis  testaceorum  Brittanniae. 

Private  publication,  London.  254  pp. 

Cox,  L.R.  1930.  The  fossil  fauna  of  the  Samana  Range  and 
some  neighbouring  areas.  Part  8,  The  Mollusca  of  the 
Hangu  shales.  Memoirs  of  the  Geological  Survey  of  In- 
dia, Palaeontologica  Indica  15:129-222. 

Cox,  L.R.,  et  al.  1969.  Systematic  descriptions.  Part  N, 
Mollusca  6,  Bivalvia,  vols.  1 and  2,  pp.  225-952  in 


68  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Moore,  R.C.,  ed.,  Treatise  on  invertebrate  paleontology. 
University  of  Kansas  Press,  Lawrence. 

Crowell,  J.C.,  and  TakeoSusuki.  1959.  Eocene  stratigraphy 
and  paleontology,  Orocopia  Mountains,  southeastern 
California.  Geological  Society  of  America,  Bulletin  70: 
581-592. 

Cushman,  J.A.  1917.  Orbitoid  Foraminifera  of  the  genus 
Orthrophragmina  from  Georgia  and  Florida.  United 
States  Geological  Survey,  Professional  Paper  108-G:  1 1 5- 
124. 

. 1920.  The  American  species  of  Orthophragmina 

and  Lepidocyclina.  United  States  Geological  Survey, 
Professional  Paper  1250:39-108. 

Cushman,  J. A.,  and  H.H.  McMasters.  1936.  Middle  Eocene 
Foraminifera  from  the  Llajas  Formation,  Ventura  Coun- 
ty, California.  Journal  of  Paleontology  1 0(6):497— 5 1 7. 

Dali,  W.H.  1886.  Reports  on  the  results  of  dredging  ...  in 
the  Gulf  of  Mexico  (1877-1878)  and  in  the  Caribbean 
Sea  (1879-1880)  ....  Report  on  the  Mollusca.  Part  1, 
Brachiopoda  and  Pelecypoda.  Harvard  Museum  of 
Comparative  Zoology,  Bulletin  12(6):  17 1-3 18. 

. 1891.  On  some  new  or  interesting  west  American 

shells  obtained  from  the  dredgings  of  the  U.  S.  Fish 
Commission  steamer  Albatross  in  1888.  United  State 
National  Museum,  Proceedings  14:173-191. 

. 1892.  Contributions  to  the  Tertiary  fauna  of  Flor- 
ida. Part  2.  Transactions  of  the  Wagner  Free  Institute 
of  Science  3:201-473. 

. 1895.  Report  on  Mollusca  and  Brachiopoda  dredged 

in  deep  water  chiefly  near  the  Hawaiian  Islands,  with 
illustrations  of  hitherto  unfigured  species  from  northwest 
America.  United  States  National  Museum,  Proceedings 
17:675-733. 

. 1902.  Illustrations  and  descriptions  of  new  unfi- 
gured, or  imperfectly  known  shells,  chiefly  American,  in 
the  U.S.  National  Museum.  United  States  National  Mu- 
seum, Proceedings  24(1264):499-566. 

. 1 903.  Synopsis  of  the  Carditacea  and  of  the  Amer- 
ican species.  Academy  of  Natural  Sciences  of  Philadel- 
phia, Proceedings  54:695-716. 

. 1904.  An  historical  and  systematic  review  of  the 

frogshells  and  tritons.  Smithsonian  Miscellaneous  Col- 
lections 47:1 14-144. 

. 1909.  Contributions  to  the  Tertiary  paleontology 

of  the  Pacific  Coast.  I.  The  Miocene  of  Astoria  and  Coos 
Bay,  Oregon.  United  States  Geological  Survey,  Profes- 
sional Paper  59:1-278,  23  pis. 

. 1915.  A monograph  of  the  molluscan  fauna  of  the 

Orthaulax  pugnax  zone  of  the  Oligocene  of  Tampa, 
Florida.  United  States  National  Museum,  Bulletin  90. 
173  pp. 

Dance,  S.P.  1976.  The  collector’s  encyclopedia  of  shells, 
2nd  ed.  McGraw-Hill  Book  Company,  New  York.  288 

pp. 

Defrance,  D.F.  1827.  Dictionnaire  des  sciences  naturelles, 
vol.  14,  Cuvier,  F.G.,  ed.  Strasbourg.  517  pp. 

Dell,  R.K.,  and  S.P.  Dance.  1963.  The  molluscan  genus 
Ranella  and  the  distribution  of  Ranella  olearium  (Lin- 


naeus). Malacoiogical  Society  of  London,  Proceedings 
35:159-167. 

Deshayes,  M.G.P.  1824-1837.  Descriptions  des  coquilles 
fossiles  des  environs  de  Paris.  2 vols.  Chez  1’auteur  and 
others,  Paris.  814  pp. 

. 1825.  Anatomic  et  monographic  du  genre  Dentale. 

Societe  d’Histoire  Naturelle  de  Paris  2:322-378. 

. 1856-1866.  Description  des  animaux  sans  ver- 

tebres  decouverts  dans  le  bassin  de  Paris.  J.-B.  Bailliere 
et  fils,  Paris.  656  pp. 

Dickerson,  R.E.  1913.  Fauna  of  the  Eocene  at  Marysville 
Buttes,  California.  University  of  California,  Publications 
in  Geological  Sciences  7(12):257-298. 

. 1914.  The  fauna  of  the  Siphonalia  sutterensis  zone 

in  the  Roseburg  Quadrangle,  Oregon.  California  Acad- 
emy of  Sciences,  Proceedings,  ser.  4,  4:1 13-128. 

. 1915.  Fauna  of  the  type  Tejon:  its  relation  to  the 

Cowlitz  phase  of  the  Tejon  Group  of  Washington.  Cal- 
ifornia Academy  of  Sciences,  Proceedings,  ser.  4,  5(3): 
33-98. 

. 1916.  Stratigraphy  and  fauna  of  the  Tejon  Eocene 

of  California.  University  of  California,  Publications  in 
Geological  Sciences  9(1 7):363— 524. 

. 1917.  Climatic  zones  of  Martinez  Eocene  time. 

California  Academy  of  Sciences,  Proceedings,  ser.  4,  7: 
193-196. 

Douville,  Henri.  1923.  Les  orbitoides  et  leur  evolution  en 
Amerique.  France  Societe  Geologie,  Bulletin,  ser.  4,  22: 
55-100. 

Dumeril,  A.M.C.  1806.  Zoologie  analytique,  ou  methode 
naturelle  de  classification  des  animaux.  Paris.  344  pp. 

Durham,  J.W.  1942.  Notes  on  Pacific  Coast  Galeodeas. 
Journal  of  Paleontology  16(2):  183-191. 

. 1950.  Cenozoic  marine  climates  of  the  Pacific  Coast. 

Geological  Society  of  America,  Bulletin  61:1243-1264. 

Edwards,  F.E.  1866.  Pyrula  angulata.  Plate  4 in  Lowry’s 
chart  of  the  characteristic  British  Tertiary  fossils. 

Eisenberg,  J.M.  1981.  A collector’s  guide  to  seashells  of  the 
world.  McGraw-Hill  Book  Company,  New  York.  240 

pp. 

Ekman,  S.  1953.  Zoogeography  of  the  sea.  Sedgwick  and 
Jackson,  London.  417  pp. 

Fagerstrom,  A.  1964.  Fossil  communities  in  paleoecology: 
their  recognition  and  significance.  Geological  Society  of 
America,  Bulletin  75:1  197-1216. 

Ferussac,  A.E.  de.  1822.  Tableaux  systematiques  des  ani- 
maux mollusques.  Paris.  1 1 1 pp. 

Filewicz,  M.V.,  and  M.E.  Hill,  III.  1983.  Calcareous  nan- 
nofossil  biostratigraphy  of  the  Santa  Susana  and  Llajas 
Formations,  north  side  Simi  Valley.  Pages  45-60  in 
Squires,  R.L.,  and  M.V.  Filewicz,  eds.,  Cenozoic  geology 
of  the  Simi  Valley  area,  southern  California.  Pacific  Sec- 
tion, Society  of  Economic  Paleontologists  and  Miner- 
alogists, Volume  and  Guidebook. 

Finlay,  H. I.  1927.  A further  commentary  on  New  Zealand 
molluscan  systematics.  Transactions  of  the  Proceedings 
of  the  New  Zealand  Institute  57:320-485. 

Finlay,  H.I.,  and  J.  Marwick.  1937.  The  Wangaloan  and 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  69 


associated  molluscan  faunas  of  Kaitangata-Green  Island 
subdivision.  New  Zealand  Geological  Survey  Branch, 
Palaeontology  Bulletin  15.  140  pp. 

Fischer,  Paul.  1 880-1 887.  Manuel  de  Conchyliologie  et  de 
Paleontologie  Conchyliologique.  F.  Savy  Librairie,  Paris. 
1369  pp. 

Fischer  von  Waldheim,  G.  1806-1807.  Museum  Demidoff, 
ou  catalogue  systematique  et  raisonne  des  curiosites  de 
la  nature  et  de  Part  ....  3 vols.  Moscou. 

Fisher,  A. G.  1966.  Spatangoids.  Part  U,  Echinodermata  3, 
pp.  543-628  in  R.C.  Moore,  ed..  Treatise  on  invertebrate 
paleontology.  University  of  Kansas  Press,  Lawrence. 

Fleming,  John.  1817-1821.  Conchology.  7 vols.  D.  Brews- 
ter’s Edinburgh  encylopaedia. 

. 1828.  A history  of  British  animals  ....  8 vols.  Bell 

and  Bradfute,  Edinburgh.  565  pp. 

Forbes,  Edward.  1838.  Malacologie  Monensis.  A catalogue 
of  the  Mollusca  inhabiting  the  Isle  of  Man  and  the  neigh- 
boring sea.  Edinburgh.  63  pp. 

Forbes,  Edward,  and  S.C.  Hanley.  1848-1853.  A history 
of  British  Mollusca  and  their  shells.  4 vols.  John  Van 
Voorst,  London.  2059  pp. 

Gabb,  W.M.  1860.  Description  of  new  species  of  American 
Tertiary  and  Cretaceous  fossils.  Academy  of  Natural 
Sciences  of  Philadelphia,  Journal,  ser.  2,  4(4):375-406. 

. 1864.  Description  of  the  Cretaceous  fossils.  Pages 

55-243,  pis.  9-32,  in  Meek,  F.B.,  and  W.M.  Gabb,  Pa- 
laentology  of  California,  Geological  Survey  of  Califor- 
nia, vol.  1,  Palaeontology.  Caxton  Press,  Philadelphia. 

. 1869.  Cretaceous  and  Tertiary  fossils.  Geological 

Survey  of  California,  vol.  2,  Palaeontology.  Caxton  Press, 
Philadelphia.  299  pp.,  36  pis. 

. 1873.  On  the  topography  and  geology  of  San  Do- 
mingo. American  Philosophical  Society,  Transactions 
15:49-259. 

Galloway,  J.J.  1 928.  A revision  of  the  family  Orbitoididae. 
Journal  of  Paleontology  2:45-69. 

Gardner,  Julia.  1926.  The  nomenclature  of  the  superspe- 
cific groups  of  Corbula  in  the  lower  Miocene  of  Florida. 
The  Nautilus  40:41-47. 

. 1 939.  Notes  on  fossils  from  the  Eocene  of  the  Gulf 

Province.  United  States  Geological  Survey,  Professional 
Paper  193-B:  17-33. 

Gardner,  Julia,  and  Edgar  Bowles.  1934.  Early  Tertiary 
species  of  gastropods  from  the  Isthmus  of  Tehuantepec. 
Journal  of  the  Washington  Academy  of  Sciences,  24(6): 
241-248. 

Givens,  C.R.  1974.  Eocene  molluscan  biostratigraphy  of 
the  Pine  Mountain  area,  Ventura  County,  California. 
University  of  California,  Publications  in  Geological  Sci- 
ences 109:1-107,  11  pis. 

. 1979.  The  gastropod  genus  Volutocristata  Gardner 

and  Bowles  (Eocene;  California,  Mexico):  a synonym  of 
Lyrischapa  Aldrich  (Eocene;  Gulf  Coast).  Tulane  Studies 
in  Geology  and  Paleontology  1 5(4):  1 17-127. 

Givens,  C.R.,  and  M.P.  Kennedy.  1976.  Middle  Eocene 
mollusks  from  northern  San  Diego  County,  California. 
Journal  of  Paleontology  50(5):954-975. 


. 1979.  Eocene  molluscan  stages  and  their  correla- 
tion, San  Diego  area,  California.  Pages  81-95  in  Abbott, 
P.L.,  ed..  Eocene  depositional  systems,  San  Diego,  Cal- 
ifornia. Pacific  Section,  Society  of  Economic  Paleonto- 
logists and  Mineralogists,  Field  Trip  Guide,  Geological 
Society  of  America  Annual  Meeting. 

Glaessner,  M.F.  1969.  Decapoda.  Part  R,  Arthropoda  4, 
pp.  400-533  in  Moore,  R.C.,  ed.,  Treatise  on  inverte- 
brate paleontology.  University  of  Kansas  Press,  Law- 
rence. 

Gmelin,  J.F.  1788-1793.  Linne’s  Systema  Naturae.  Editio 
13.  3 vols. 

Goldfuss,  A.  1826-1833.  Petrefacta  Germaniae.  Amz, 
Diisseldorf.  252  pp. 

Grabau,  A.W.  1904.  Phylogeny  of  Fusus  and  its  allies. 
Smithsonian  Miscellaneous  Collections  44(14 17):  1-192. 

Grant,  U.S.,  IV,  and  H.R.  Gale.  1931.  Catalogue  of  the 
marine  Pliocene  and  Pleistocene  Mollusca  of  California. 
San  Diego  Society  of  Natural  History,  Memoirs  1:1- 
1036. 

Grant,  U.S.,  IV,  and  L.G.  Hertlein.  1938.  The  West  Amer- 
ican Cenozoic  Echinoidea.  University  of  California, 
Publications  in  Mathematical  and  Physical  Sciences  2: 
1-226. 

Grasse,  P.P.  1968.  Mollusques  gasteropodes  et  scapho- 
podes.  Traite  de  Zoologie  Anatomie,  Systematique, 
Biologie.  Libraires  de  L’Academie  de  Medecine,  Paris. 
Vol.  5,  1083  pp. 

Gray,  J.E.  1808-1856.  Synopsis  of  the  contents  of  the  Brit- 
ish Museum.  British  Museum,  London.  370  pp. 

. 1847.  A list  of  the  genera  of  Recent  Mollusca,  their 

synonyms  and  types.  Zoological  Society  of  London,  Pro- 
ceedings 15:129-219. 

Gregorio,  M. A.  de.  1890.  Monographic  de  la  faune  Eoce- 
nique  de  l’Alabama  et  surtout  de  ceile  de  Claiborne  de 
l’etage  Parisien.  Annales  de  Geologie  et  de  Paleontologie, 
livraisons  7 et  8.  316  pp. 

Habe,  Tadashige.  1968.  Shells  of  the  western  Pacific  in 
color,  vol.  2,  revised  edition.  Hoikusha  Publishing  Com- 
pany, Osaka.  233  pp. 

Hanna,  G.D.  1924.  Rectifications  of  nomenclature.  Cali- 
fornia Academy  of  Sciences,  Proceedings,  ser.  4,  13(10): 
151-186. 

Hanna,  G.D.,  and  L.G.  Hertlein.  1943.  Characteristic  fos- 
sils of  California.  California  Division  of  Mines  and  Ge- 
ology, Bulletin  1 18(2):  165-1 82. 

Hanna,  M.A.  1 927.  An  Eocene  invertebrate  fauna  from  the 
La  Jolla  Quadrangle,  California.  University  of  Califor- 
nia, Publications  in  Geological  Sciences  1 6(8):247— 398. 

Harris,  G.G.  1897.  Catalogue  of  Tertiary  Mollusca  in  the 
Department  of  Geology,  British  Museum  (Natural  His- 
tory), Part  1.  The  Australasian  Tertiary  Mollusca.  Lon- 
don. 407  pp. 

Heckel,  P.H.  1972.  Recognition  of  ancient  shallow  marine 
environments.  Pages  226-286  in  Rigby,  J.K.,  and  W.K. 
Hamblin,  eds.,  Recognition  of  ancient  sedimentary  en- 
vironments. Society  of  Economic  Paleontologists  and 
Mineralogists,  Special  Publication  16. 


70  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Herrmannsen,  A.N.  1846-1852.  Indicis  generum  mala- 
cozoorum  primordia.  2 vols.  Theodore  Fischeri,  Cassel. 
1494  pp. 

Hertlein,  L.G.  1933.  A new  gryphaeoid  oyster  from  the 
Eocene  of  California.  San  Diego  Society  of  Natural  His- 
tory, Transactions  7:277-282. 

Hertlein,  L.G.,  and  U.S.  Grant,  IV.  1944.  The  Cenozoic 
Brachiopoda  of  western  North  America.  University  of 
California,  Publications  in  Mathematical  and  Physical 
Sciences  3: 1-236. 

. 1972.  The  geology  and  paleontology  of  the  marine 

Pliocene  of  San  Diego,  California  (Paleontology:  Pele- 
cypoda).  San  Diego  Society  of  Natural  History,  Memoir 
2,  pt.  2B.  409  pp.,  57  pis. 

Hickman,  CJ  S.  1969.  The  Oligocene  marine  molluscan 
fauna  of  the  Eugene  Formation  in  Oregon.  University 
of  Oregon  Museum  of  Natural  History,  Bulletin  16:1  — 
112. 

. 1980.  Paleogene  marine  gastropods  of  the  Keasey 

Formation  in  Oregon.  Bulletins  of  American  Paleon- 
tology 78: 1-1 12. 

Hinds,  R.B.  1 843.  Descriptions  of  new  shells  from  the  col- 
lection of  Captain  Sir  Edward  Belcher  . . . during  the 
years  1836-1842  ....  Zoological  Society  of  London, 
Proceedings  11:36-49. 

Houbrick,  R.S.  1977.  Reevaluation  and  new  description  of 
the  genus  Bittium  (Cerithiidae).  The  Veliger  20(2):  101  — 
106. 

Howell,  B.F.  1962.  Worms.  Part  W,  Miscellanea,  pp.  1 44— 
177  in  Moore,  R.C.,  ed.,  Treatise  on  invertebrate  pa- 
leontology. University  of  Kansas  Press,  Lawrence. 

Hyatt,  Alpheus.  1894.  Phylogeny  of  an  acquired  charac- 
teristic. American  Philosophical  Society,  Proceedings  32: 
349-647. 

Ingram,  W.M.  1940.  A new  Gisortia.  Journal  of  the  Wash- 
ington Academy  of  Sciences  30(9):376— 377. 

. 1942.  Type  fossil  Cypraeidae  of  North  America. 

Bulletins  of  American  Paleontology  27(1 04):  1 —32. 

International  Commission  on  Zoological  Nomenclature.  1964. 
International  code  of  zoological  nomenclature.  Inter- 
national Trust  for  Zoological  Nomenclature,  London. 
176  pp. 

Iredale,  T.  1913.  The  generic  name  to  be  used  for  Murex 
tritonis  Linne.  The  Nautilus  27:55-56. 

. 1916.  On  some  new  and  old  molluscan  names. 

Malacological  Society  of  London,  Proceedings  12:27- 
37. 

Jeletzky,  J.A.  1966.  Comparative  morphology,  phylogeny, 
and  classification  of  fossil  Coleoidea.  The  University  of 
Kansas  Paleontological  Contributions,  Mollusca,  article 
7,  pp.  1-162. 

Jousseaume,  F.P.  1887.  La  famille  des  Cancellariidae.  Le 
Naturaliste,  annales  9,  ser.  2,  pp.  155-157,  163,  1 92— 
194,  213,  214,  221-223. 

Jukes-Browne,  A.J.  1908.  On  the  genera  of  Veneridae  rep- 
resented in  the  Cretaceous  and  older  Tertiary  deposits. 
Malacological  Society  of  London,  Proceedings  8:148- 
177. 


Jung,  Peter.  1974.  A revision  of  the  Family  Seraphsidae 
(Gastropoda:Strombacea).  Palaeontographica  Ameri- 
cana 8(47):5— 72. 

Keen,  A.  M.  1944.  Catalogue  and  revision  of  the  gastropod 
subfamily  Typhinae.  Journal  of  Paleontology  18(1 ): 50— 
72. 

. 1951.  The  molluscan  names  in  Reiner’s  “Tavole.” 

The  Nautilus  65(1 ): 8—  1 5 . 

. 1971.  Sea  shells  of  tropical  West  America,  2nd  ed. 

Stanford  University  Press,  Stanford.  1064  pp. 

Keen,  A.M.,  and  Herdis  Bentson.  1944.  Check  list  of  Cal- 
ifornia Tertiary  marine  Mollusca.  Geological  Society  of 
America,  Special  Papers  56:1-280. 

Keen,  A.M.,  and  Eugene  Coan.  1974.  Marine  molluscan 
genera  of  western  North  America,  2nd  ed.  Stanford  Uni- 
versity Press,  Stanford.  208  pp. 

Keenan,  M.F.  1932.  The  Eocene  Sierra  Blanca  limestone 
at  the  type  locality  in  Santa  Barbara  County,  California. 
Transactions  of  the  San  Diego  Society  of  Natural  History 
7:53-84. 

Kennett,  J.P.  1982.  Marine  geology.  Prentice-Hall,  Engle- 
wood Cliffs,  New  Jersey.  813  pp. 

Kennett,  J.P.,  et  al.  1975.  Cenozoic  paleoceanography  in 
the  southwest  Pacific  Ocean,  Antarctic  glaciation,  and 
the  development  of  the  cirum-Antarctic  current.  Pages 
1 155-1 169  in  Kennett,  J.P.,  et  al.,  eds.,  Initial  reports 
of  the  Deep  Sea  Drilling  Project,  vol.  29.  United  States 
Government  Printing  Office,  Washington,  D.C. 

Kew,  W.S.W.  1919.  Structure  and  oil  resources  of  the  Simi 
Valley,  southern  California.  United  States  Geological 
Survey,  Bulletin  691:323-347. 

. 1920.  Cretaceous  and  Cenozoic  Echinoidea  of  the 

Pacific  Coast  region  of  North  America.  University  of 
California,  Publications  in  Geological  Sciences  1 2:23— 
236. 

. 1924.  Geology  and  oil  resources  of  a part  of  Los 

Angeles  and  Ventura  Counties,  California.  United  States 
Geological  Survey,  Bulletin  753:1-202. 

Kira,  Tetsuaki.  1965.  Shells  of  the  western  Pacific  in  color, 
vol.  1,  2nd  ed.  Hoikusha  Publishing  Company,  Osaka. 
224  pp. 

Kleinpell,  R.M.,  and  D.W.  Weaver.  1963.  Oligocene  bio- 
stratigraphy of  the  Santa  Barbara  embayment,  Califor- 
nia. University  of  California,  Publications  in  Geological 
Sciences  43: 1-250. 

Kobelt,  Wilhelm.  1883.  Iconographie  der  schalentragenden 
europaischen  Meeresconchilien.  Heft  3.  Wiesbaden. 

Laiming,  Boris.  1940a.  Some  foraminiferal  correlations  in 
the  Eocene  of  San  Joaquin  Valley,  California.  Sixth  Pa- 
cific Science  Congress,  Proceedings  2:535-568. 

. 1 940b.  Foraminiferal  correlations  in  Eocene  of  San 

Joaquin  Valley,  California.  American  Association  of  Pe- 
troleum Geologists,  Bulletin  24:1923-1939. 

. 1943.  Eocene  foraminiferal  correlations  in  Cali- 
fornia. California  Division  of  Mines,  Bulletin  1 18:193- 
198. 

Lamarck,  J.B.  de.  1799.  Prodome  d’une  nouvelle  classifi- 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  71 


cation  des  coquilles  ....  Memoires  de  la  Societe  d’His- 
toire  naturelle  de  Paris,  pp.  63-91. 

. 1801.  Systeme  des  animaux  sans  vertebres  ou  ta- 
bleau general  des  classes,  des  ordres,  et  des  genres  de  ces 
animaux.  Paris.  432  pp. 

. 1802-1809.  Memoires  sur  les  fossiles  des  environs 

de  Paris.  Annales  du  Museum  National  d’Histoire  Na- 
turelle, vols.  1-9,  12,  14.  Variously  paged. 

. 1811.  Suite  de  la  determination  des  especes  de 

mollusques  testaces:  continuation  du  genre  Volute  et 
Mitre.  Annales  du  Museum  d’Histoire  Naturelle  Paris 
17:54-80,  195-222. 

. 1815-1822.  Histoire  naturelle  des  animaux  sans 

vertebres.  7 vols.  Paris. 

Lambert,  Jules.  1902.  Description  des  echinides  fossiles  de 
la  province  de  Barcelone.  Memoires  de  la  Societe  Geo- 
logique  de  France,  Paleontologie,  Memoire  24:1-128. 

Latreille,  P.A.  1825.  Families  naturelles  du  regne  animal, 
exposees  succintement  et  dans  un  order  analytique,  avec 
l’indication  de  leurs  genres.  Second  edition.  Paris. 

Lea,  Isaac.  1833.  Contributions  to  geology.  Carey,  Lea,  and 
Blanchard,  Philadelphia.  227  pp. 

Leach,  W.E.  1819.  A list  of  invertebrate  animals  discovered 
by  H.M.S.  ship  Isabella,  in  a voyage  to  the  arctic  regions. 
Appendix  II,  pp.  61-64  in  Ross,  C.J.,  Voyage  of  dis- 
covery . . . exploring  Baffin’s  Bay.  Appendix  I,  pp.  1- 
252. 

Leriche,  Maurice.  1906.  Contribution  a l’etude  des  Pois- 
sons fossiles  du  Nord  de  la  France  et  des  regions  voisines. 
Memoire  Societe  Geologie  Nord  5:1-430. 

Lewis,  W.D.  1940.  The  geology  of  the  upper  Las  Llajas 
Canyon  area,  Santa  Susana  Mountains,  California.  Cal- 
ifornia Institute  of  Technology,  Unpublished  Master  of 
Science  thesis.  74  pp. 

Lindner,  Gert.  1978.  Seashells  of  the  world.  Van  Nostrand 
Reinhold  Company,  New  York.  271  pp. 

Link,  H.F.  1807.  Beschreibung  der  Naturalien-Sammlung 
der  Universitat  zu  Rostok,  vol.  1.  Variously  paged. 

Linne,  Carl  von.  1758.  Systema  naturae  per  regna  tria  na- 
turae, Editio  decima,  reformata,  Regnum  animale,  vol. 

1 . Holmiae.  824  pp. 

. 1766-1767.  Systema  naturae  per  regna  tria  naturae, 

Editio  duodecima,  reformata,  Regnum  animale,  vol.  1 . 
Holmiae.  1327  pp. 

Ludbrook,  N.H.  1960.  Scaphopoda.  Part  I,  Mollusca  1,  pp. 
37-41  in  Moore,  R.C.,  ed..  Treatise  on  invertebrate  pa- 
leontology. University  of  Kansas  Press,  Lawrence. 

MacLeay,  W.S.  1838.  On  the  brachyurous  decapod  Crus- 
tacea brought  from  the  Cape  by  Dr.  Smith.  Vol.  5,  In- 
vertebratae,  pp.  53-71,  in  Andrew  Smith,  M.D.,  Illus- 
trations of  the  zoology  of  South  Africa. 

Mallory,  V.S.  1959.  Lower  Tertiary  biostratigraphy  of  the 
California  Coast  Ranges.  American  Association  of  Pe- 
troleum Geologists,  Tulsa.  416  pp. 

Marincovich,  Louie,  Jr.  1977.  Cenozoic  Naticidae  (Mol- 
lusca:Gastropoda)  of  the  northeastern  Pacific.  Bulletins 
of  American  Paleontology  70(294):  169-494. 


Marshall,  W.B.  1917.  The  Wangaloa  beds.  New  Zealand 
Institute,  Transactions  49(38):450-460,  4 pis. 

Mawe,  John.  1823.  The  Linnaean  system  of  conchology. 
London.  207  pp. 

May er-Ey mar,  Karl.  1877.  Systematisches  Verzeichnissder 
Versteinerungen  des  Parisian  der  Umgegend  von  Ein- 
siedeln.  Beitragen  Geologisch  Karte  Schweiz  14(26):  100 

pp. 

McLaughlin,  R.P.,  and  C. A.  Waring.  1915.  Petroleum  in- 
dustry of  California.  California  Mining  Bureau  Bulletin 
69.  519  pp.  Atlas. 

McMasters,  J.H.  1932.  The  Eocene  Llajas  Formation,  Ven- 
tura County,  California.  Stanford  University,  Unpub- 
lished Master  of  Arts  thesis.  70  pp. 

McMillan,  N.F.  1968.  British  shells.  Frederick  Wame  and 
Company,  London.  196  pp. 

Meek,  F.B.  1876.  A report  on  the  invertebrate  Cretaceous 
and  Tertiary  fossils  of  the  upper  Missouri  country.  United 
States  Geological  Survey,  Territory  Annual  Report  9:1- 
629,  45  pis. 

Merriam,  C.W.  1941.  Fossil  Turritellas  from  the  Pacific 
Coast  region  of  North  America.  University  of  California, 
Publications  in  Geological  Sciences  26(1):  1-2 14. 
Merriam,  C.W. , and  F.E.  Turner.  1937.  The  Capay  middle 
Eocene  of  northern  California.  University  of  California, 
Publications  in  Geological  Sciences  24(6):9 1 — 1 14. 
Miles,  G.A.  1981.  Planktonic  foraminifers  of  the  lower 
Tertiary  Roseburg,  Lookingglass,  and  Flournoy  For- 
mations (Umpqua  Group),  southwest  Oregon.  Pages  85- 
103  in  Armentrout,  J.M.,  ed..  Pacific  Northwest  Ceno- 
zoic biostratigraphy.  Geological  Society  of  America, 
Special  Paper  184. 

Miller,  A.K.  1947.  Tertiary  nautiloids  of  the  Americas. 

Geological  Society  of  America,  Memoir  23:1-234. 
Milne-Edwards,  Henri.  1852.  Observations  sur  les  affinites 
zoologiques  et  la  classification  naturelle  des  crustaces. 
Annales  Science  Naturelle,  Zoologie,  ser.  3,  18:109-166. 
Milne-Edwards,  Henri,  and  Jules  Haime.  1848-1 850a.  Re- 
cherches  sur  les  polypes.  Annales  Science  Naturelle  de 
Paris,  vols.  9-13,  15-16. 

. 1850b.  A monograph  of  the  British  fossil  corals. 

Introduction;  corals  from  the  Tertiary  and  Cretaceous 
formations.  Palaeontolographical  Society,  London.  85 

pp. 

Montagu,  George.  1 803-1 808.  Testacea  Britannica,  or  nat- 
ural history  of  British  shells,  marine,  land  and  fresh- 
water. 2 vols.  London.  789  pp. 

Monterosato,  T.A.  di.  1 884.  Nomenclatura  generica  e spe- 
zifica  di  alcune  conghiglie  mediterranee.  Palermo.  152 

pp. 

Montfort,  P.D.  1808-1810.  Conchyliologie  systematique 
et  classification  methodique  de  coquilles.  2 vols.  Paris. 
676  pp. 

Moore,  E.J.  1983.  Tertiary  marine  pelecypods  of  Califor- 
nia and  Baja  California:  Nuculidae  through  Malleidae. 
United  States  Geological  Survey,  Professional  Paper 
1228-A:A1-A108,  27  pis. 


72  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


Morch,  O.A.L.  1852-1853.  Catalogus  conchyliorum  quae 
reliquit  D.  Alphonso  d’Aguiira  et  Gadea  Comes  de  Yol- 
di.  8 vols.  Hafniae. 

. 1858.  Note surle genre  Volutharpa Fischer.  Journal 

de  Conchyliologie,  sen  2,  3:40-44. 

Morris,  P.A.  1966.  Pacific  Coast  shells,  2nd  ed.  Houghton 
Mifflin  Company,  Boston.  297  pp. 

. 1975.  Shells  of  the  Atlantic  and  Gulf  Coasts  and 

the  West  Indies,  3rd  ed.  Houghton  Mifflin  Company, 
Boston.  330  pp. 

Muir-Wood,  H.M.,  G.F.  Elliott,  and  Kotora  Hatai.  1965. 
Mesozoic  and  Cenozoic  Terebratellidina.  Part  H,  Brach- 
iopoda,  vol.  2,  pp.  8 1 6-857  in  Moore,  R.C.,  ed..  Treatise 
on  invertebrate  paleontology.  University  of  Kansas  Press, 
Lawrence. 

Muller,  O.F.  1776.  Zoologiae  Danicae  prodromus  seu  an- 
imalium  Daniae  et  Norvegiae  indigenarum  characteres, 
nomina  et  synonyma  imprimis  popuiarium,  8 vols.  Hav- 
niae.  282  pp. 

Muller,  Johannes,  and  F.G.J.  Henle.  1837.  Ueber  die  Gat- 
tungen  der  Plagiostomen.  Archiv  Naturgeschichte  3:394- 
401,  434. 

. 1838.  On  the  generic  characters  of  cartilaginous 

fishes,  with  descriptions  of  new  genera.  Charlesworth’s 
Magazine  of  Natural  History  2:33-37,  88-91. 

Munier-Chalmas,  E.C.P.A.  1864.  Societe  Linnaeus  Nor- 
mandie, Bulletin  8:99,  108. 

Nelson,  R.N.  1925.  A contribution  to  the  paleontology  of 
the  Martinez  Eocene  of  California.  University  of  Cali- 
fornia, Publications  in  Geological  Sciences  15(1  1 ): 397— 
466. 

Newton,  R.B.  1922.  Eocene  Mollusca  from  Nigeria.  Geo- 
logical Survey  of  Nigeria,  Bulletin  3:1-140. 

Nilsen,  T.H.  1973.  Facies  relations  in  the  Eocene  Tejon 
Formation  of  the  San  Emigdio  and  western  Tehachapi 
Mountains,  California.  Pages  7-23  in  Fischer,  Pete,  ed., 
Sedimentary  facies  changes  in  Tertiary  rocks,  California 
Transverse  and  southern  Coast  Ranges.  Society  of  Eco- 
nomic Paleontologists  and  Mineralogists,  Los  Angeles. 

Nomland,  J.O.  1916.  Corals  from  the  Cretaceous  and  Ter- 
tiary of  California  and  Oregon.  University  of  California, 
Publications  in  Geological  Sciences  9(5):59-76. 

Nordsieck,  Fritz.  1968.  Die  europaischen  Meeres-gehause- 
schnecken  (Prosobranchia).  Gustav  Fischer  Verlag, 
Stuttgart.  273  pp. 

. 1969.  Die  europaischen  Meeresmuscheln  (Bival- 

via).  Gustav  Fischer  Verlag,  Stuttgart.  256  pp. 

Okada,  Hisatake,  and  David  Bukry.  1980.  Supplementary 
modification  and  introduction  of  code  numbers  to  the 
low-latitude  coccolith  biostratigraphic  zonation  (Bukry, 
1973;  1975).  Marine  Micropaleontology  5:321-325. 

Olsson,  A.  A.  1961.  Mollusks  of  the  tropical  eastern  Pacific: 
Panama-Pacific  Pelecypoda.  Paleontological  Research 
Institution,  Ithaca.  574  pp.,  86  pis. 

Orbigny,  A.D.  d\  1843.  Paleontologie  franyaise.  Terrains 
cretaces  3:1-807. 


Orr,  W.N.,  and  M.A.  Kooser.  1971.  Oregon  Eocene  deca- 
pod Crustacea.  The  Ore  Bin  33:1 19-129. 

Palmer,  D.B.K.  1923.  A fauna  from  the  middle  Eocene 
shales  near  Vacaville,  California.  University  of  Califor- 
nia, Publications  in  Geological  Sciences  1 4(8):289— 3 1 8 . 

Palmer,  K.V.W.  1927-1929.  The  Veneridae  of  eastern 
America,  Cenozoic  and  Recent.  Palaeontographica 
Americana  1 (5):209— 522,  45  pis. 

. 1967.  A comparison  of  certain  Eocene  molluscs  of 

the  Americas  with  those  of  the  western  Tethys.  Pages 
183-193  in  Adams,  C.G.,  and  D.V.  Ager,  eds..  Aspects 
of  Tethyan  biogeography.  Systematics  Association  Pub- 
lication, no.  7. 

. 1974.  Composition  with  relationships  of  Paleocene 

and  Eocene  molluscan  fauna  of  the  East  Americas.  Pages 
468-485  in  Contributions  to  the  geology  and  paleo- 
biology of  the  Caribbean  and  adjacent  areas.  Verhand- 
lungen  der  Naturforschenden  Gesellschaft  in  Basel,  vol. 
84. 

Palmer,  K.V.W. , and  D.C.  Brann.  1965-1966.  Catalogue 
of  the  Paleocene  and  Eocene  Mollusca  of  the  southern 
and  eastern  United  States.  Bulletins  of  American  Pa- 
leontology 48:1-1057  (2  parts). 

Perry,  George.  1810-181 1.  Arcana,  or  the  Museum  of  Nat- 
ural History;  containing  the  most  recent  discovered  ob- 
jects ....  London. 

Peterson,  G.L.,  and  P.L.  Abbott.  1979.  Mid-Eocene  cli- 
matic change,  southwestern  California  and  northwestern 
Baja  California.  Palaeogeography,  Palaeoclimatology, 
Palaeoecology  26:73-87. 

Philippi,  R.A.  1853.  Handbuch  der  Conchyliologie  und 
Malakozoologie.  Halle.  547  pp. 

Pilsbry,  H.A.,  and  A.  A.  Olsson.  1954.  Systems  of  the  Volu- 
tidae.  Bulletins  of  American  Paleontology  35(1 5 2):27 1 — 
306. 

Poli,  G.S.  1791-1795.  Testacea  utriusque  Siciliae,  eorum- 
que  historia  et  anatomia.  3 vols.  Parma. 

Poore,  R.Z.  1980.  Age  and  correlation  of  California  Paleo- 
gene benthic  foraminiferal  stages.  United  States  Geo- 
logical Survey,  Professional  Paper  1 1 62-C.  8 pp. 

Powell,  A. W.B.  1942.  The  New  Zealand  Recent  and  fossil 
Mollusca  of  the  family  Turridae,  with  general  notes  on 
turrid  nomenclature  and  systematics.  Bulletin  of  Auck- 
land Institute  and  Museum,  no.  2.  188  pp. 

Quayle,  E.H.  1932.  Fossil  corals  of  the  genus  Turbinolia 
from  the  Eocene  of  California.  Transactions  of  the  San 
Diego  Society  of  Natural  History  7(1 0):9 1 — 1 09. 

Rafinesque,  C.S.  1 809.  Indice  d’ittiologia  Siciliana.  Mes- 
sina, pp.  12-167. 

. 1815.  Analyse  de  la  nature,  ou  tableau  de  Funivers 

et  des  corps  organisees.  Palermo. 

Rathbun,  M.J.  1 926.  The  fossil  stalk-eyed  Crustacea  of  the 
Pacific  slope  of  North  America.  United  States  National 
Museum,  Bulletin  138:1-155. 

Rehder,  H.A.  1973.  The  family  Harpidae  of  the  world. 
Indo-Pacific  Mollusca  3(16):207-274. 

Reuss,  A.E.  1859.  Zur  Kenntnis  fossiler  Krabben.  Aka- 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  73 


demie  Wissenschaftlich  Wien,  Denkschrift  17:1-90,  24 
pis. 

Rios,  E.C.  1970.  Coastal  Brazilian  seashells.  Museu  Ocea- 
nografico  de  Rio  Grande.  255  pp. 

Risso,  J.A.  1810-1826.  Histoire  naturelle  des  principales 
productions  de  1’Europe  meridionale  et  particulierement 
de  celles  des  environs  de  Nice  et  des  Alpes  maritimes. 
5 vols.  Paris. 

Roding,  P.F.  1798.  Museum  Boltenianum  sive  catalogus 
cimeliorum  e tribus  regnis  naturae  quae  olim  collegerat. 
J.F.  Bolten,  M.D.,  ed.,  Pars  Secunda.  Typis  Johan  Chris- 
ti  Trappii,  Hamburg.  199  pp. 

Romer,  Eduard.  1857.  Kritische  Untersuchung  der  Arten 
des  Molluskengeschlechts  Venus  bei  Linne  und  Gmelin 
mit  Beriicksichtigung  der  spater  Beschriebenen  Arten.  J. 
Georg  Luckhardt,  Cassel.  136  pp. 

Ross,  C. A.  1974.  Paleogeography  and  provinciality.  Pages 
1-17  in  Ross,  C.A.,  ed.,  Paleogeographic  provinces  and 
provinciality.  Society  of  Economic  Paleontologists  and 
Mineralogists,  Tulsa. 

Sacco,  Frederico.  1890-1904.  I Molluschi  dei  terreni  ter- 
ziari  del  Piemonte  e della  Liguria  descritti  da  Luigi  Bel- 
lardi.  Musei  di  Zoologia  ed  Anatomia  comparata  della 
R.  Universita  di  Torino,  Bollettino  pts.  6-30. 

Saul,  L.R.  1983.  Notes  on  Paleogene  turritellas,  venericar- 
dias,  and  molluscan  stages  of  the  Simi  Valley  area,  Cal- 
ifornia. Pages  71-80  in  Squires,  R.L.,  and  M.V.  Filewicz, 
eds.,  Cenozoic  geology  of  the  Simi  Valley  area,  southern 
California.  Pacific  Section,  Society  of  Economic  Pale- 
ontologists and  Mineralogists,  Volume  and  Guidebook. 

Savin,  S.M.,  R.G.  Douglas,  and  F.C.  Stehli.  1975.  Tertiary 
marine  paleotemperatures.  Geological  Society  of  Amer- 
ica, Bulletin  86:1499-1510. 

Schenck,  H.G.  1926.  Cassididae  of  western  America.  Uni- 
versity of  California,  Publications  in  Geological  Sciences 
16(4):69-98. 

. 1929.  Discocyclina  in  California.  Transactions  of 

the  San  Diego  Society  of  Natural  History  5(1 4):2 1 1-240. 

. 1931.  Cephalopods  of  the  genus  Aturia  from  west- 
ern North  America.  University  of  California,  Publica- 
tions in  Geological  Sciences  1 9(  1 9):435— 490. 

. 1936.  Nuculid  bivalves  of  the  genus  Acila.  Geo- 
logical Society  of  America,  Special  Paper  4:1-149. 

Schmidt,  F.C.  1818.  Versuch  fiber  die  beste  Einrichtung 
zur  Aufstellung,  Behandlung  und  Aufbewahrung  der  ver- 
schieden  Naturkorper  und  Gegenstande  der  Kunst. 
Gotha.  252  pp. 

Schumacher,  C.F.  1817.  Essais  d'un  nouveau  systeme  des 
habitations  des  vers  testaces.  4 vols.  Cophenhagen.  287 

pp. 

Schymiczek,  H.B.  1983a.  Benthic  Foraminifera  and  paleo- 
bathymetry  of  the  Eocene  Llajas  Formation,  north  side 
of  Simi  Valley,  California.  Pages  97-108  in  Squires,  R.L., 
and  M.V.  Filewicz,  eds.,  Cenozoic  geology  of  the  Simi 
Valley  area,  southern  California.  Pacific  Section,  Society 
of  Economic  Paleontologists  and  Mineralogists,  Volume 
and  Guidebook. 

. 1 983b.  Benthic  Foraminifera  and  paleobathymetry 


of  the  Eocene  Llajas  Formation,  southwestern  Santa  Su- 
sana  Mountains,  California.  California  State  University, 
Northridge,  Master  of  Science  thesis.  76  pp. 

Schymiczek,  H.B. , and  R.L.  Squires.  1981.  Biostratigraphy 
of  discocyclinid-bearing  beds.  Eocene  Llajas  Formation, 
southwestern  Santa  Susana  Mountains,  California. 
American  Association  of  Petroleum  Geologists,  Bulletin 
65:989. 

Scopoli,  J.A.  1777.  Introductio  ad  historiam  naturaleum 
sistens  genera  lapidum,  plantarum  et  animalus  ....  Pra- 
gae.  506  pp. 

Seiden,  Hyman.  1972.  Geology  of  Las  Llajas  Canyon,  Ven- 
tura County,  California.  University  of  California  Los 
Angeles,  Unpublished  Master  of  Science  thesis.  33  pp. 

Shepard,  F.P.,  R.R.  Lankford,  and  E.D.  Milow.  1957.  La 
Jolla  area  field  trip  syllabus.  Pacific  Section,  Society  of 
Economic  Paleontologists  and  Mineralogists,  Field  Trip 
Guide.  10  pp. 

Smith,  J.P.  1919.  Climatic  relations  of  the  Tertiary  and 
Quaternary  faunas  of  the  California  region.  California 
Academy  of  Sciences,  Proceedings,  ser.  4,  9(4):  1 23— 173. 

Smith,  J.T.  1970.  Taxonomy,  distribution,  and  phylogeny 
of  the  cymatiid  gastropods  Argobuccinum,  Fusitriton, 
Mediargo,  and  Priene.  Bulletins  of  American  Paleon- 
tology 56:445-573. 

. 1975.  Age,  correlation,  and  possible  Tethyan  affin- 
ities of  mollusks  from  the  Lodo  Formation  of  Fresno 
County,  California.  Pages  464-483  in  Weaver,  D.W., 
G.R.  Homaday,  and  Ann  Tipton,  eds.,  Future  energy 
horizons  of  the  Pacific  Coast;  Paleogene  symposium  and 
selected  technical  papers.  American  Association  of  Pe- 
troleum Geologists,  Society  of  Economic  Paleontologists 
and  Mineralogists,  and  Society  of  Economic  Geologists, 
Pacific  Sections,  Annual  Meeting. 

Solander,  D.C.  1766.  Description  of  species.  Pp.  1-43  in 
Gustavo  Brander,  Fossilia  Hantoniensia  collecta,  et  in 
Museuo  Britannico  deposita.  London. 

Sowerby,  James.  1 8 1 2a- 1 846a.  Mineral  conchology,  7 vols. 
London. 

. 1837b-1844b.  Mineral-conchologie  Grossbritan- 

niens  ....  Deutsch  bearbeitet  von  E.  Desor.  Durchge- 
sehen  und  mit  Anmerkungen  und  Berichtigungen  ver- 
sehen  von  L.  Agassiz.  Solothum.  689  pp. 

Spengler,  Lorenz.  1794-1802.  Over  det  taskallede  slaegt 
tellineme.  5 vols.  Skriv.  Naturhist.  Selskabet.  Copen- 
hagen. 

Squires,  R.L.  1977.  Middle  Eocene  molluscan  assemblage 
and  stratigraphy,  lower  Piru  Creek,  Transverse  Ranges, 
California.  California  Division  of  Mines  and  Geology, 
Special  Report  129:81-86. 

. 1979.  New  macrofossil  occurrences,  Stewart  bed, 

middle  Eocene  Llajas  Formation,  Llajas  Canyon,  Santa 
Susana  Mountains,  California.  Geological  Society  of 
America,  Cordilleran  Section  Meeting  [Abstracts]  1 1(3): 
129. 

. 1981.  A transitional  alluvial  to  marine  sequence: 

the  Eocene  Llajas  Formation,  southern  California.  Jour- 
nal of  Sedimentary  Petrology  5 1 (3):923— 938. 


74  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


. 1983a.  New  mollusks  from  the  lower  middle  Eo- 
cene Llajas  Formation,  southern  California.  Journal  of 
Paleontology  57:354-362. 

. 1983b.  Eocene  Llajas  Formation,  Simi  Valley, 

southern  California.  Pages  81-95  in  Squires,  R.L.,  and 
M.V.  Filewicz,  eds.,  Cenozoic  geology  of  the  Simi  Valley 
area,  southern  California.  Pacific  Section,  Society  of  Eco- 
nomic Paleontologists  and  Mineralogists,  Volume  and 
Guidebook. 

Stanton,  T.W.  1896.  The  faunal  relations  of  the  Eocene  and 
Upper  Cretaceous  on  the  Pacific  Coast.  United  States 
Geological  Survey,  Annual  Report  17:1005-1060. 

Stewart,  R.B.  1927.  Gabb’s  California  fossil  type  gastro- 
pods. Academy  of  Natural  Sciences  of  Philadelphia,  Pro- 
ceedings 78:287-447,  pis.  20-32. 

. 1930.  Gabb’s  California  Cretaceous  and  Tertiary 

type  lamellibranchs.  Academy  of  Natural  Sciences  of 
Philadelphia,  Special  Publication  3:1-314,  17  pis. 

. 1946.  Geology  of  Reef  Ridge  Coalinga  district  Cal- 
ifornia. United  States  Geological  Survey,  Professional 
Paper  205C:8 1-1 15,  pis.  11-17. 

Stipp,  T.F.  1 943.  Simi  oil  field.  Pages  4 1 5-424  in  Geologic 
formations  and  economic  development  of  the  oil  and 
gas  fields  of  California.  California  Division  of  Mines, 
Bulletin  1 1 8,  pt.  3. 

Stoliczka,  Ferdinand.  1868.  The  Gastropoda  of  the  Cre- 
taceous rocks  of  southern  India.  Memoirs  of  the  Geo- 
logical Survey  of  India,  Palaeontologica  Indica,  ser.  5, 
2:1-498. 

. 1870-1871.  Cretaceous  fauna  of  southern  India. 

The  Pelecypoda,  with  a review  of  all  known  genera  of 
this  class,  fossil  and  Recent.  Geological  Survey  of  India, 
Palaeontologica  Indica,  ser.  6,  3:1-535. 

Suter,  Henry.  1913.  Manual  of  the  New  Zealand  Mollusca. 
Government  Printer,  Wellington.  1 1 20  pp. 

Sutherland,  J. A.  1 966.  A new  species  of  Architectonica  from 
the  Santa  Susana  Mountains,  Ventura  County,  Califor- 
nia. Los  Angeles  County  Museum,  Contributions  in  Sci- 
ence 1 1 7: 1-4. 

Swainson,  William.  1820-1833.  Zoological  illustrations,  2 
ser.  London. 

. 1 840.  A treatise  on  malacology  or  shells  and  shell- 
fish. London.  419  pp. 

Tebble,  Norman.  1976.  British  bivalve  seashells,  2nd  ed. 
Her  Majesty’s  Stationery  Office,  Edinburgh.  212  pp. 

Tegland,  N.M.  1933.  The  fauna  of  the  type  Blakeley  upper 
Oligocene  of  Washington.  University  of  California,  Pub- 
lications in  Geological  Sciences  23:81-174. 

Thiele,  Johannes.  1925.  Gastropoden  der  deutschen  Tief- 
see-expedition  II.  Wissenschaftlich  Ergebnis  Deutsch 
Tiefsee-Expedition  1 7(2):36— 282. 

. 1929-1935.  Handbuch  der  systematischen  Weich- 

tierkunde.  2 vols.  Jena.  1 134  pp. 

Troschel,  F.H.  1852.  Verzeichniss  der  durch  Herm  Dr.  v. 
Tschudi  in  Peru  gesammelten  Conchylien.  Archiv  fur 
Naturgeschichte  18(1):  15 1-208. 

Turner, F.E.  1938.  Stratigraphy  and  Mollusca  of  the  Eocene 


of  western  Oregon.  Geological  Society  of  America,  Spe- 
cial Paper  10: 1-1 30. 

Vail,  P.R.,  R.M.  Mitchum,  Jr.,  and  S.  Thompson,  III.  1977. 
Seismic  stratigraphy  and  global  changes  of  sea  level,  part 
4:  Global  cycles  of  relative  changes  of  sea  level.  Pages 
83-97  in  Payton,  C.E.,  ed..  Seismic  stratigraphy  — ap- 
plications to  hydrocarbon  exploration.  American  As- 
sociation of  Petroleum  Geologists,  Tulsa,  Oklahoma. 

Vaughan,  T.W.  1900.  Eocene  and  lower  Oligocene  coral 
faunas  of  the  United  States  with  a few  doubtfully  Cre- 
taceous species.  United  States  Geological  Survey,  Mono- 
graph 39:1-263. 

. 1936.  New  species  of  orbitoidal  Foraminifera  of 

the  genus  Discocyclina  from  the  lower  Eocene  of  Ala- 
bama. Journal  of  Paleontology  10:255-256. 

. 1945.  American  Paleocene  and  Eocene  larger  Fo- 
raminifera. Geological  Society  of  America,  Memoir  9(  1 ): 
1-175. 

Verastegui,  Pedro.  1953.  The  Pelecypod  genus  Venericar- 
dia  in  the  Paleocene  and  Eocene  of  western  North  Amer- 
ica. Palaeontographica  Americana  3(25):  1-1 12. 

Vokes,  H.E.  1935.  Notes  on  the  variation  and  synonymy 
of  Ostrea  idriaensis  Gabb.  University  of  California,  Pub- 
lications in  Geological  Sciences  23(9):29 1-304. 

. 1937.  The  gastropod  genus  Harpa  in  the  Eocene 

of  the  western  United  States.  Journal  of  Paleontology 
1 1(1):  10—12,  pi.  2. 

. 1939.  Molluscan  faunas  of  the  Domengine  and 

Arroyo  Hondo  Formations  of  the  California  Eocene. 
Annals  of  the  New  York  Academy  of  Sciences  38: 1-246. 

. 1940.  Paleoecology  of  the  fauna  of  the  Domengine 

Formation,  middle  Eocene,  California.  Sixth  Pacific  Sci- 
ence Congress,  Proceedings  2:597-605. 

. 1980.  Genera  of  the  Bivalvia:  a systematic  and 

bibliographic  catalogue  (revised  and  updated).  Paleon- 
tological Research  Institution,  Ithaca.  307  pp. 

Waring,  C.  A.  1914.  Eocene  horizons  of  California.  Journal 
of  Geology  22(8):782-785. 

. 1917.  Stratigraphic  and  faunal  relations  of  the  Mar- 
tinez to  the  Chico  and  Tejon  of  southern  California. 
California  Academy  of  Sciences,  Proceedings,  ser.  4,  7(4): 
41-124. 

Weaver,  C.E.  1905.  Contribution  to  the  palaeontology  of 
the  Martinez  Group.  University  of  California,  Publi- 
cations in  Geological  Sciences  4(5):  101-123. 

. 1912.  A preliminary  report  on  the  Tertiary  pa- 
leontology of  western  Washington.  Washington  Geolog- 
ical Survey,  Bulletin  15:1-80,  15  pis. 

. 1943.  Paleontology  of  the  marine  Tertiary  for- 
mations of  Oregon  and  Washington.  University  of 
Washington,  Publications  in  Geology  5(1-3):  1-789. 

. 1953.  Eocene  and  Paleocene  deposits  at  Martinez, 

California.  University  of  Washington  Publications  in 
Geology  7: 1-102. 

Weaver,  C.E.,  and  K.V.W.  Palmer.  1922.  Fauna  from  the 
Eocene  of  Washington.  University  of  Washington  Pub- 
lications in  Geology  1(3):  1-56. 

Weaver,  C.E.,  et  al.  1944.  Correlation  of  the  marine  Ce- 


Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks  75 


nozoic  formations  of  western  North  America.  Geolog- 
ical Society  of  America,  Bulletin  55:569-598. 

Weaver,  C.S.,  and  duPont,  J.E.  1970.  Living  volutes.  Del- 
aware Museum  of  Natural  History,  Monograph  Series 
No.  1,  Greenville,  Delaware.  375  pp. 

Wells,  J.W.  1956.  Scleractinia.  Part  F,  Coelenterata,  pp. 
328-444  in  Moore,  R.C.,  ed.,  Treatise  on  invertebrate 
paleontology.  University  of  Kansas  Press,  Lawrence. 

. 1957.  Corals.  Pages  1087-1 104  in  Hedgpeth,  J.W. , 

ed..  Treatise  on  marine  ecology  and  paleoecology.  Geo- 
logical Society  of  America,  Memoir  67,  vol.  1. 

Wenz,  Wilhelm.  1938-1944.  Gastropoda.  Vol.  6,  7 parts, 
pp.  1-1639  in  Schindewolf,  O.H.,  ed.,  Handbuch  der 
Palaozoologie.  Gebriider  Borntraeger,  Berlin. 

White,  C.A.  1889.  On  invertebrate  fossils  from  the  Pacific 
Coast.  United  States  Geological  Survey,  Bulletin  51:1— 
102. 

Wolfe,  J.A.  1 977.  Paleogene  floras  from  the  Gulf  of  Alaska 
region.  United  States  Geological  Survey,  Professional 
Paper  997.  108  pp. 

Wood,  William.  1818-1828.  Index  testaceologicus.  8 vols. 
Privately  published,  London.  188  pp. 

Woodring,  W.P.  1925.  Miocene  Mollusca  from  Bowden 
Jamaica,  pelecypods  and  scaphopods.  Carnegie  Institute 
of  Washington,  Publication  366:1-564,  40  pis. 

. 1928.  Miocene  mollusks  from  Bowden,  Jamaica. 


Part  2,  Gastropods  and  discussion  of  results.  Carnegie 
Institute  of  Washington,  Publication  385:1-564. 

. 1966.  The  Panama  land  bridge  as  a sea  barrier. 

American  Philosophical  Society,  Proceedings  1 10:425- 
433. 

Woodward,  S.P.  1851-1856.  A manual  of  the  Mollusca:  or 
rudimentary  treatise  of  recent  and  fossil  shells.  John 
Weale,  London.  330  pp. 

Zinsmeister,  W.J.  1974.  Paleocene  biostratigraphy  of  the 
Simi  Hills,  Ventura  County,  California.  University  of 
California,  Riverside,  Unpublished  Ph.D.  dissertation. 
236  pp. 

. 1982.  Late  Cretaceous-early  Tertiary  molluscan 

biogeography  of  the  southern  circum-Pacific.  Journal  of 
Paleontology  56:84-102. 

. 1983a.  Late  Paleocene  (“Martinez  Provincial 

Stage”)  molluscan  fauna  from  the  Simi  Hills,  Ventura 
County,  California.  Pages  61-70  in  Squires,  R.L.,  and 
M.V.  Filewicz,  eds.,  Cenozoic  geology  of  the  Simi  Valley 
area,  southern  California.  Pacific  Section,  Society  of  Eco- 
nomic Paleontologists  and  Mineralogists,  Volume  and 
Guidebook. 

. 1 983b.  New  late  Paleocene  molluscs  from  the  Simi 

Hills,  Ventura  County,  California.  Journal  of  Paleon- 
tology 57:1282-1303. 

Accepted  for  publication  7 February  1984. 


76  Contributions  in  Science,  Number  350 


Squires:  Simi  Valley  Eocene  Mollusks 


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MOLLUSCAN  PALEONTOLOGY  AND  BIOSTRATIGRAPHY  OF  THE 
LOWER  MIOCENE  UPPER  PART  OF  THE  LINCOLN  CREEK 
FORMATION  IN  SOUTHW  ESTERN  W ASHINGTON 

Ellen  J.  Moore 


Contributions  in  Science,  Number  351 
Natural  History  Museum  of  Los  Angeles  County 
13  July  1984 


Vessels  loading  lumber  at  Knappton,  Washington,  in  1908.  When  it  was  founded  in  1867,  Knappton  was  called  Cementville, 
and  concretionary  rocks  served  as  the  raw  material  for  the  cement.  (Photograph  courtesy  of  Dale  Estoos  and  David  Lee  Myers.) 


ISSN  0459-8113 

Natural  History  Museum  of  Los  Angeles  County 
900  Exposition  Boulevard 
Los  Angeles,  California  90007 


PREFACE 


Knappton,  Washington,  on  the  Columbia  River  north  of  Astoria,  Oregon,  was  named  for 
Jabez  Burrell  Knapp,  who  built  a cement  plant  there  in  1867,  calling  the  town  Cementville. 
The  plant  included  a large  kiln  and  a cement-barrel  factory,  but  was  unable  to  continue 
production  for  more  than  a few  years  because  of  lack  of  sufficient  raw  material:  fossiliferous 
concretions  and  concretionary  layers.  Knapp  then  built  a sawmill,  which  supported  the 
company  town  for  70  years.  In  1941,  the  planing  mill  caught  fire  and  the  blaze  quickly 
spread  to  other  buildings.  The  wooden  water  main  burned  through,  leaving  the  fire  fighters 
helpless,  and  the  town  was  destroyed.  Only  a monument  and  pilings  remain  at  the  site  to 
reflect  its  past  history. 

A few  fossil  mollusks  were  collected  from  the  Knappton  area  in  the  early  1900’s  and 
deposited  in  the  National  Museum  of  Natural  History  at  Washington,  D.C.,  and  the  Cali- 
fornia Academy  of  Sciences  at  San  Francisco.  The  small  collections  contain  usually  poorly 
preserved  material  from  both  the  upper  part  of  the  Lincoln  Creek  Formation  and  the  over- 
lying  Astoria  Formation,  which  were  not  differentiated  at  that  time. 

Controversy  over  the  age  of  the  Astoria  Formation,  whose  type  locality  is  just  across  the 
river,  began  in  1865  and  revolved  especially  around  the  presence  of  the  fossil  cephalopod 
Aturia.  This  cephalopod  also  was  known  to  occur  in  the  upper  part  of  the  Lincoln  Creek 
Formation  on  the  Washington  side  of  the  Columbia  River,  and  casual  references  to  Knappton 
occasionally  appeared  in  the  scientific  literature.  It  was  not  until  1979,  however,  that  a 
detailed  geologic  map  of  the  Knappton  area  was  prepared  by  Ray  E.  Wells  and  issued  by 
the  U.S.  Geological  Survey. 

The  first  major  collections  of  fossils  from  Knappton  were  made  by  James  L.  Goedert  and 
his  wife,  Gail.  The  presence  of  vertebrate  fossils,  chiefly  whales,  at  Knappton  led  Goedert 
to  contact  the  Los  Angeles  County  Museum  of  Natural  History  in  1979.  His  offer  to  donate 
the  collections  to  the  Los  Angeles  Museum,  and  the  recognition  of  the  importance  of  the 
invertebrate  fossils  in  those  collections  by  Edward  C.  Wilson,  Curator  of  Invertebrate  Pa- 
leontology, has  led  to  publications  on  the  barnacles  by  Victor  A.  Zullo,  the  sponges  by  J. 
Keith  Rigby  and  D.E.  Jenkins,  and  the  mollusks  described  herein. 

This  paper  is  dedicated  to  James  L.  Goedert  whose  generosity  has  made  it  possible  for 
specialists  to  study  the  remarkable  Knappton  fossils. 


CONTENTS 


ABSTRACT  1 

INTRODUCTION  1 

PREVIOUS  WORK  2 

CHRONOSTRATIGRAPHY 3 

PALEOECOLOGY 6 

MOLLUSCAN  PALEONTOLOGY  8 

Abbreviations 9 

Gastropods 9 

Trochidae 9 

Epitoniidae 10 

Naticidae 10 

Cassididae 1 0 

Buccinidae? 12 

Neptuneidae  12 

Fusinidae 16 

Volutidae 16 

Turridae 24 

Acteonidae 25 

Pelecypods 25 

Nuculidae 25 

Nuculanidae 25 

Solemyidae 25 

Limopsidae 25 

Mytilidae 26 

I.imidae  26 

Ostreidae 28 

Lucinidae 28 

Thyasiridae  28 

Cardiidae 30 

Tellinidae 30 

Periplomatidae 30 

Teredinidae 30 

Scaphopod 34 

Dentaliidae  34 

Cephalopods 34 

Aturidae 34 

Sepiidae?  34 

FOSSILS  OTHER  THAN  MOLLUSKS 34 

Sponges 34 

Corals 36 

Brachiopod  36 

Echinoids 36 

LOCALITIES 36 

ACKNOWLEDGMENTS  39 

LITERATURE  CITED 39 


MOLLUSCAN  PALEONTOLOGY  AND  BIOSTRATIGRAPHY  OF  THE 
LOWER  MIOCENE  UPPER  PART  OF  THE  LINCOLN  CREEK 
FORMATION  IN  SOUTHWESTERN  WASHINGTON 

Ellen  J.  Moore1 


ABSTRACT.  The  uppermost  part  of  the  Lincoln  Creek  Formation 
in  the  Knappton  area  of  southwestern  Washington  is  assigned  to  the 
lower  Miocene  upper  part  of  the  Juanian  Molluscan  Stage  (=Sauce- 
sian  Foraminiferal  Stage)  and  the  Liracassis  apta  Molluscan  Zone. 
This  part  of  the  Lincoln  Creek  Formation  is  correlated  with  the 
upper  part  of  the  Pysht  Formation  of  the  Twin  River  Group  in 
northwestern  Washington. 

Fossils,  invertebrates  and  vertebrates,  are  preserved  in  concretions 
that  erode  out  of  landslides  and  accumulate  as  float  along  the  Co- 
lumbia River.  The  molluscan  fauna  of  the  upper  part  of  the  Lincoln 
Creek  Formation  consists  of  33  species  in  30  genera,  including  five 
newly  described  species  in  the  genera  “ Bathybembix ,”  Ancistrolepis, 
Musashia,  Modiolus,  and  Acesta.  Eastern  Pacific  species  in  the  genus 
Musashia  are  reviewed.  The  fauna  lived  at  depths  between  100  and 
350  m,  a bathymetric  range  substantiated  by  the  inferred  ranges  of 
16  associated  phyla  also  preserved  as  fossils  at  Knappton.  The  nau- 
tiloid  cephalopod  Aturia,  which  is  common  in  the  collections,  in- 
dicates that  the  water  temperature  may  have  been  as  high  as  1 6°C 
at  a depth  of  100  m.  The  abundance  of  preserved  organic  material 
suggests  that  free  oxygen  was  depleted  in  the  sediment  below  the 
level  of  bioturbation. 

INTRODUCTION 

Fossiliferous  concretions  that  have  eroded  out  of  landslide 
blocks  from  the  upper  part  of  the  Lincoln  Creek  Formation 
occur  as  float  along  the  Columbia  River  near  the  site  of 
Knappton  in  southwestern  Washington  (Figs.  1, 2).  The  Lin- 
coln Creek  Formation  in  this  area  is  part  of  a homocline, 
and  the  sequence  dips  eastward.  The  upper  part  consists  of 
poorly  bedded,  locally  laminated,  bioturbated,  concretionary 
dark-gray  siltstone.  The  formation  is  unconformable  upon 
the  upper  Eocene  siltstones  of  Cliff  Point  unit  (Wells,  1979). 

The  invertebrate  fauna  described  in  this  report  is  from  the 
uppermost  part  of  the  Lincoln  Creek  Formation  (LAM  Loc. 
5842)  and  is  early  Miocene  in  age.  The  fauna  is  assigned  to 
the  upper  part  of  the  Juanian  Molluscan  Stage  (=Saucesian 
Foraminiferal  Stage)  and  the  Liracassis  apta  [Echinophoria 
apta]  Molluscan  Zone.  The  Lincoln  Creek  Formation  is  over- 
lain  by  the  lower  Miocene  part  of  the  Astoria  Formation, 

Contributions  in  Science,  Number  351,  pp.  1-42 
Natural  History  Museum  of  Los  Angeles  County,  1984 


assigned  to  the  Pillarian  Molluscan  Stage  and  the  Vertipecten 
fucanus  Molluscan  Zone  (Fig.  3). 

Although  the  fossiliferous  concretions  are  collected  as  float 
on  the  bank  of  the  Columbia  River,  an  approximate  stratig- 
raphy is  preserved  in  the  landslides  because  wedge-shaped 
blocks  move  south  toward  the  river  bank  parallel  with  the 
strike  of  the  rocks.  One  locality,  informally  called  the  “glass 
sponge  bed”  (LAM  Loc.  5852),  and  others  below  it,  called 
the  “decapod  crustacean  bed”  (LAM  Loc.  5843)  and  the 
“gooseneck  barnacle  bed”  (LAM  Loc.  5844),  all  lie  strati- 
graphically  below  the  major  mollusk-bearing  unit  (the  Aturia 
bed)  described  here  (LAM  Loc.  5842).  Faunas  in  these  four 
beds  are  segregated  in  the  float  in  proper  stratigraphic  po- 
sition. The  lack  of  mixing  is  also  characteristic  of  faunas 
typical  of  the  overlying  Astoria  Formation  (LAM  Loc.  5863). 

The  geology  in  the  area  is  complicated  by  landslides,  faults, 
by  few  road  cuts,  and  by  vegetative  cover  (Figs.  4,  5).  Gen- 
erally, only  sections  100  m or  less  can  be  measured  in  tidal 
exposures,  and  the  relationship  between  exposures  is  often 
difficult  to  discern  (Figs.  6,  7).  Mapping  by  Wells  (1979) 
portrays  the  complexities  in  the  area. 

Despite  the  poor  exposures,  some  interpretations  can  be 
made.  The  concretions  are  continuously  being  reworked  from 
the  landslides  and  new  accumulations  appear  with  sufficient 
regularity  to  be  collected  every  two  weeks  at  low  tide.  The 
concretions  therefore  are  randomly  distributed  throughout 
the  unit. 

Most  of  the  concretions  are  spherical  and  composed  of 
fine-grained  siltstone  with  calcareous  cement.  A few  are  ce- 
mented by  quartz  in  the  central  part  and  calcite  at  the  rim. 
I believe  that  the  concretions  formed  early  in  diagenesis, 
because  the  mollusks  preserved  in  the  concretions  are  almost 
always  complete  specimens  that  are  neither  broken  nor 


1 . U.S.  Geological  Survey,  345  Middlefield  Road  MS-9 1 5,  Menlo 
Park,  California  94025,  and  Research  Associate,  Invertebrate  Pa- 
leontology Section,  Natural  History  Museum  of  Los  Angeles  County. 


ISSN  0459-8113 


124°  123° 


Figure  1.  Index  map  of  the  Pacific  coast  showing  localities  men- 
tioned in  the  text. 


crushed.  Delicate  features  of  snails  such  as  the  long,  narrow 
siphonal  canal  of  Priscofusus  and  the  T-shaped  axial  ribs  of 
Ancistrolepis  are  preserved  intact.  Some  of  the  concretions 


do  not  have  any  obvious  organic  center,  but  most  contain 
fossils,  many  are  bioturbated,  and  most  contain  fecal  pellets. 

Of  particular  interest  as  an  aid  to  diagenetic  interpretation 
is  the  preservation  of  complex  sequences  of  laminated  calcite, 
sparry  calcite,  barite,  and  quartz,  precipitated  in  that  order, 
within  the  phragmocone  chambers  of  the  cephalopod  Aturia. 
This  sequential  mineralization  is  currently  being  studied,  but 
it  is  obvious  now  that  the  laminated  calcite  was  precipitated 
first  followed  by  sparry  calcite,  barite,  and  quartz. 

PREVIOUS  WORK 

James  L.  Goedert,  in  collaboration  with  Gail  H.  Goedert, 
began  collecting  fossils  in  the  Knappton  area  in  the  late  1970’s 
and  donated  the  collections  to  the  Natural  History  Museum 
of  Los  Angeles  County.  The  first  invertebrate  collections  were 
made  over  a relatively  large  area  and  assigned  the  locality 
number  LAM  5787.  Later  observations  led  Goedert  to  realize 
that  the  concretions  and  other  fossiliferous  material  on  the 
terrace  represented  separate  faunal  zones.  Thus  he  separated 
subsequent  collections  into  three  localities:  the  lowest  (LAM 
Loc.  5844)  contains  abundant  gooseneck  barnacles  ( Arco - 
scalpellum)  and  the  trace  fossil  Tisoa;  the  middle  (LAM  Loc. 
5843)  contains  many  decapod  crustaceans;  and  the  upper- 
most (LAM  Loc.  5842)  contains  siliceous  sponges,  mollusks 
including  large  specimens  of  the  cephalopod  Aturia,  and 
abundant  marine  vertebrates.  Still  later,  he  was  able  to  sep- 
arate the  siliceous  sponge-bearing  locality  (LAM  Loc.  5852) 
from  the  other  three  localities.  At  that  time,  he  divided  the 
collecting  area  into  four  informal  faunal  units.  Beginning  at 
the  base  of  the  section  these  are;  Unit  I (LAM  Loc.  5844), 
Unit  II  (LAM  Loc.  5843),  Unit  III  (LAM  Loc.  5852),  and 
Unit  IV  (LAM  Loc.  5842)  (Fig.  3). 

Victor  A.  Zullo  (1982)  described  the  barnacles  from  Units 
I and  IV.  From  Unit  I,  he  described  two  species  of  gooseneck 
barnacles,  Arcoscalpellum  knapptonensis  and  A.  raricosta- 
tum,  and  assigned  the  unit  to  the  upper  Eocene.  From  Unit 
IV,  Zullo  described  the  archaeobalinid  Solidoba/anus  (Hes- 
peribalanus)  sp.  alf.  S.  (//.)  sookensis  (Cornwall)  and  assigned 
the  unit  to  the  upper  Oligocene. 

J.  Keith  Rigby  and  David  E.  Jenkins  (1983)  described 
sponges  from  Units  II,  III,  and  IV.  Eurete  goederti  was  de- 
scribed from  Unit  III  and  Aphrocallistes  polytretos  was  de- 
scribed from  Units  II,  III,  and  IV.  Rigby  and  Jenkins  also 
identified  A.  polytretos  in  three  other  places:  a limestone 
quarry  in  the  Bear  River  area  northwest  of  Knappton  (LAM 
Loc.  5802)  that  is  in  the  upper  Eocene  siltstones  of  CliffPoint 
unit  (Wells,  1979);  in  the  type  area  of  the  lower  and  middle 
Miocene  Astoria  Formation  at  Astoria,  Oregon;  and  in  the 
upper  Oligocene  and  lower  Miocene  Yaquina  Formation, 
south  of  Newport,  Oregon.  Thus  E.  goederti  is  early  Miocene 
in  age  and  the  range  of  A.  polytretos  is  late  Eocene  to  middle 
Miocene. 

J.  Dale  Nations,  Northern  Arizona  University,  is  studying 
the  decapod  crustaceans  from  Units  II  and  IV,  Carole  S. 
Hickman,  University  of  California,  is  studying  micromol- 
lusks  from  the  section  at  Knappton,  and  Bruce  J.  Welton, 
Chevron  Oil  Field  Research  Company,  the  fish.  Birds  and 
marine  mammals,  collected  throughout  the  section  but  most 


2 Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks 


123°48'  123°46’ 


Formation,  Ta  = Astoria  Formation,  Tb  = Tertiary  basaltic  sill,  and  Qls  = Quartemary  landslide  material. 


commonly  from  Unit  IV,  are  in  the  vertebrate  collections  of 
the  Natural  History  Museum  of  Los  Angeles  County. 

CHRONOSTRATIGRAPHY 

Molluscan  stages  were  proposed  for  the  Pacific  northwest 
Tertiary  section  by  Addicott  (1976c)  and  by  Armentrout 
(1975,  1977),  and  molluscan  zones  by  Durham  (1944),  with 
revisions  and  additions  by  both  Addicott  (1976c)  and  Ar- 
mentrout (1977).  Subsequent  work  (Allison,  1978;  Marin- 
covich,  1979;  Moore,  1984)  has  demonstrated  the  usefulness 
of  these  stages  and  zones  (Fig.  3). 

Stratigraphic  sections  representing  the  major  Oligocene  part 
of  the  Juanian  Molluscan  Stage  have  been  extensively  studied 
and  their  molluscan  faunas  described  and  illustrated  (Teg- 
land,  1933;  Weaver,  1942;  Durham,  1944;  Armentrout,  1973; 
Addicott,  1976a,  1976b). 

The  upper  part  of  the  Lincoln  Creek  Formation  is  assigned 


to  the  upper  part  of  the  Juanian  Molluscan  Stage,  equivalent 
to  the  upper  part  of  the  Liracassis  apta  Molluscan  Zone  (Fig. 
3).  The  upper  part  of  the  formation  is  of  early  Miocene  age 
and  equivalent  in  age  to  the  earliest  part  of  the  Saucesian 
Foraminiferal  Stage. 

The  upper  part  of  the  Lincoln  Creek  Formation  in  south- 
western Washington,  assigned  to  the  part  of  the  Juanian  that 
is  of  late  Oligocene  age,  contains  a molluscan  fauna  that 
remained  essentially  the  same  in  species  composition 
throughout  the  late  Oligocene.  The  strata  exposed  near 
Knappton,  which  represent  the  highest  exposed  part  of  the 
Lincoln  Creek  Formation  and  the  part  of  the  Juanian  Mol- 
luscan Stage  that  is  of  earliest  Miocene  age,  contains  five  new 
species,  representing  about  1 5%  of  the  molluscan  fauna  in 
the  unit.  I interpret  this  as  indicating  that  this  part  of  the 
section  is  rarely  preserved  and  that  the  fauna  is  transitional 
between  the  well-known  part  (upper  Oligocene)  of  the  Jua- 
nian and  the  Pillarian  (lower  Miocene).  The  fauna  has  more 


Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks  3 


Figure  3.  Stratigraphic  position  of  fossil  localities  in  the  Lincoln  Creek  Formation  and  adjacent  formations  in  southwestern  Washington. 


species  in  common  with  the  Juanian  than  with  the  Pillarian, 
which  may  indicate  that  it  is  closer  in  age  to  the  Juanian  or 
lived  at  depths  more  commonly  represented  in  the  Juanian. 
The  mollusks  from  the  upper  part  of  the  Lincoln  Creek 


Formation  are  similar  enough  to  the  molluscan  fauna  from 
the  upper  part  of  the  Pysht  Formation  of  the  Twin  River 
Group,  exposed  along  the  Strait  of  Juan  de  Fuca,  to  suggest 
a partial  correlation  of  those  formations.  Addicott  (1976b: 


4 Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks 


Figure  4.  View  looking  southwest  toward  Knappton  monument  at 
upper  left  and  past  fossil  localities  along  the  coast  at  the  right,  [m 
monument,  f = fossil  localities.] 

442)  considered  the  molluscan  fauna  in  the  upper  part  of  the 
Pysht  Formation  to  belong  to  the  Liracassis  apta  Molluscan 
Zone.  Although  L.  apta  was  not  collected  from  the  Pysht 
Formation,  Addicott  believed  that  this  was  due  to  that  unit’s 
relatively  shallow-water  depositional  environment  rather  than 
to  the  organism’s  extinction.  On  the  basis  of  foraminiferal 
evidence,  Addicott  assigned  the  upper  part  of  the  Pysht  For- 
mation to  the  Saucesian.  “ Bathybembix”  hickmanae  n.  sp., 
Bruclarkia  yaquinana,  and  Megasurcula  sp.  cf.  M.  wyno- 
ocheensis  suggest  correlation  of  the  upper  part  of  the  Lincoln 
Creek  Formation  with  the  upper  part  of  the  Pysht  Formation. 

Musashia  ( Nipponomelon ) shikamai  n.  sp.  and  Bruclarkia 
yaquinana,  from  the  upper  part  of  the  Lincoln  Creek  For- 
mation at  Knappton,  were  previously  reported  from  strata 


Figure  5.  Landslide  exposure  of  upper  part  of  the  Lincoln  Creek 
Formation  along  the  Columbia  River. 


Figure  6.  Tidal  exposure  of  upper  part  of  the  Lincoln  Creek  For- 
mation on  the  Columbia  River  terrace. 

no  older  than  the  Clallam  Formation  (Addicott,  1976c:  14- 
15)  of  early  Miocene  age,  equivalent  to  the  Pillarian  Mol- 
luscan Stage.  The  occurrence  of  these  species  in  the  upper 
part  of  the  Lincoln  Creek  Formation  extends  their  range 


Figure  7.  Concretionary  layer  in  upper  part  of  the  Lincoln  Creek 
Formation  exposed  in  a landslide  block  along  the  Columbia  River. 


Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks  5 


Depth  in  meters 

o 1000 2000 3000 4000 

Gastropods 

Bathybembix 

Sthenorytis 

Ancistrolepis 

Musashia  (Musashia)  • • • • 

Musashia  (Nipponomelon)  • 

Aforia 

Pelecypods 

Acila  (Acila) 

Portlandia  (Portlandia)  * • 

Acharax 

Limopsis 

Modiolus 

Acesta  ( Plicacesta ) 

Lucmoma 

Thyasira  (Conchocele)  • • • 

Cochlodesma 

Cephalopod 

Aturia,  based  on  Nautilus  • • 


Figure  8.  Depth  range  of  living  species  of  genera  of  fossils  found  in  concretions  in  the  Aturia  bed  near  the  top  of  the  Lincoln  Creek  Formation, 
at  Knappton,  Washington.  The  overlapping  ranges  indicate  a depth  for  the  assemblage  of  about  100  to  350  meters,  equivalent  to  the  outer 
continental  shelf  or  the  upper  continental  slope. 


downward  into  the  upper  part  of  the  Juanian  Molluscan  Stage. 
This  further  refines  the  Juanian  Molluscan  Stage  and  indi- 
cates an  interval  in  the  earliest  part  of  the  Saucesian  when  a 
single  molluscan  province  may  have  extended  from  south- 
western to  northwestern  Washington. 

The  generic  composition  of  the  Knappton  fauna  (“ Bath - 
ybembix Liracassis,  Ancistrolepis,  and  Musashia ) is  similar 
to  other  assemblages  of  mollusks  collected  from  the  Lincoln 
Creek  Formation  and  its  correlative  units,  and  the  conditions 
of  deposition  seem  also  to  be  similar. 

Musashia  ( Nipponomelon ) weaveri  survived  throughout  the 
entire  Oligocene.  Musashia  (N.)  shikamai  n.  sp.,  also  present 
in  the  Clallam  Formation  (Addicott,  1976c,  pi.  3,  fig.  27) 
and  in  the  upper  part  of  the  Poul  Creek  Formation,  replaced 
M.  weaveri  in  the  upper  part  of  the  Lincoln  Creek  Formation. 

Liracassis  durhami  Kanno,  which  occurs  with  Liracassis 
apta  at  Knappton  (Moore,  1984),  was  present  during  the 
upper  part  of  the  Juanian  Stage,  then  became  locally  extinct, 
but  survived  through  the  Pillarian  Stage  in  the  upper  part  of 
the  Poul  Creek  Formation  in  the  Gulf  of  Alaska.  Although 
common  in  the  collections  from  Knappton,  L.  durhami  is 
rare  in  other  exposures  of  the  Lincoln  Creek  Formation. 

Ancistrolepis  clarki  teglandae  occurs  in  the  Oligocene  part 
of  the  Juanian  Stage  and  was  preceded  by  A.  clarki  clarki  in 
the  Matlockian  Stage.  Ancistrolepis  jimgoederti  n.  sp.  occurs 


in  the  upper  part  of  the  Lincoln  Creek  at  Knappton,  in  the 
lower  Miocene  part  of  the  Juanian  Stage. 

The  uppermost  part  of  the  Lincoln  Creek  Formation  at 
Knappton  is  assigned  to  the  earliest  Miocene  on  the  basis  of 
its  molluscan  fauna.  Two  fossil  localities  of  Goedert,  units  3 
and  4 are  of  early  Miocene  age;  units  1 and  2 may  be  of  late 
Oligocene  age.  The  upper  part  of  the  Lincoln  Creek  For- 
mation at  Knappton  is  correlative  with  the  upper  part  (lower 
Miocene)  of  the  Pysht  Formation.  The  placement  of  the  up- 
per part  of  the  Lincoln  Creek  Formation  in  the  lower  Miocene 
corroborates  the  assignment  of  the  upper  part  of  the  Juanian 
Stage  to  the  lower  Miocene  (Addicott,  1976b:442). 

PALEOECOLOGY 

The  Lincoln  Creek  Formation  in  southwestern  Washington 
represents  the  accumulation  of  sediment  in  a relatively  stable 
basin  environment  that  shallowed  to  the  southeast.  The  mol- 
luscan fauna  of  the  lowermost  Miocene  part  of  the  Lincoln 
Creek  Formation  at  Knappton  lived  at  dephs  between  100 
and  350  m (Fig.  8). 

The  mollusk-bearing  concretions  are  highly  bioturbated 
with  numerous  burrows.  Preserved  fecal  pellets  occur  as  small 
spherical  ooids  within  the  body  chambers  of  gastropods, 
elongate  pelloids  within  the  septal  chambers  of  the  cepha- 


6 Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks 


lopod  Aturia  (Figs.  156,  157),  and  both  types  scattered 
throughout  the  concretions.  Seventeen  phyla  are  represented 
in  the  fauna,  and  they  indicate  a healthy  community  of  or- 
ganisms living  in  water  of  normal  salinity.  The  abundance 
of  Aturia  suggests  that  the  water  temperature  may  have  been 
as  high  as  16°C  with  shallow  embayments  nearby  with  tem- 
peratures as  high  as  24°C  for  egg  laying  (Cochram,  Rye,  and 
Landman,  1981:477).  The  abundance  of  preserved  organic 
material  suggests  that  the  sediment  below  the  level  of  bio- 
turbation  was  dysaerobic,  reduced  in  oxygen  (0. 1-0.5  ml/ 
liter),  and  that  regular  sedimentation  gradually  buried  the 
organic  material  without  destroying  it  or  the  organisms  that 
lived  there. 

Zullo  (1982  and  written  commun.,  1982)  described  Soli- 
dobalanus  (Hesperibalanus)  aff.  S.  (H.)  sookensis  (Cornwall) 
from  the  upper  part  of  the  Lincoln  Creek  Formation  at 
Knappton.  Although  this  archeobalanid  barnacle  is  repre- 
sentative of  a sessile-benthic  group  usually  found  at  subtidal 
to  inner-shelf  depths,  the  basal  plates  of  the  Knappton  spec- 
imens suggest  attachment  to  shells  or  wood,  which  could 
have  been  transported  to  the  site  of  deposition. 

Rigby  and  Jenkins  (1983)  described  and  assigned  sponges 
from  the  upper  part  of  the  Lincoln  Creek  Formation  to  the 
genera  Aphrocallistes  and  Eurete;  the  distribution  of  living 
species  of  both  genera  suggests  that  they  lived  at  a depth  of 
approximately  300  to  350  m. 

The  foraminifers  in  the  upper  part  of  the  Lincoln  Creek 
Formation  in  the  Grays  River  quadrangle,  just  east  of 
Knappton,  indicate  water  depths  of  300  to  900  m (Rau  in 
Wolfe  and  McKee,  1972:42). 

James  C.  Ingle,  Jr.  (written  commun.,  1982)  examined  thin 
sections  made  from  the  centers  of  mollusk-bearing  concre- 
tions. He  concluded:  “All  of  the  evidence  in  your  thin  sec- 
tions suggests  deposition  occurred  on  a continental  slope  or 
marginal  basin  associated  with  impingement  of  the  oxygen 
minimum  layer  producing  anaerobic  or  dysaerobic  condi- 
tions ....  The  core  of  this  oxygen  depleted  water  mass  com- 
monly occurs  at  a depth  between  200  and  600  m off  Cali- 
fornia today  and  the  foraminifera  identified  in  your  thin 
sections  support  this  depth  range  ( Globulimina , Epistomi- 
nella,  Bolivina,  Uvigerina).  There  is  evidence  of  redeposition 
of  some  of  the  material  from  shallower  environments  with 
neritic-littoral  echinoderm  spines  and  thick  walled  porcela- 
neous  foraminifera  present  in  several  samples.  In  addition, 
the  rare  glauconite  fragments  were  likely  redeposited  from 
the  adjacent  shelf-edge  or  outer  neritic  area.” 

A sample  of  sediment  from  the  head  of  the  active  landslide 
at  the  west  end  of  the  major  fossil-bearing  concretion  locality 
(LAM  5842)  and  samples  from  a measured  section  strati- 
graphically  below  have  yielded  Zemmorian  age  foraminiferal 
assemblages  and  suggest,  as  a conservative  estimate,  that  the 
water  depth  was  midslope,  1000  m or  possibly  deeper  (Kris- 
tin McDougall,  written  commun.,  1982;  James  C.  Ingle,  Jr., 
written  commun.,  1983). 

Bruce  J.  Welton,  studying  the  fish  remains  from  Knappton, 
has  found  bones,  scales,  and  teeth  to  be  abundant  at  all  the 
localities  and  bony-fish  otoliths  to  be  common  in  almost  all 
samples.  Preliminary  identifications  show  that  the  fish  in- 


clude seven  genera  of  sharks  representing  six  families,  and 
one  family  of  bony  fish,  in  addition  to  numerous  unidentified 
otoliths  and  isolated  bones.  According  to  Welton  (written 
commun.,  1982), 

Ecologically,  the  sharks  are  represented  by  two  epipe- 
lagic  genera  ( Cetorhinus  and  Eugomphodus)  and  five 
genera  ( Centrophorus , Chlamydoselachus,  Notorynchus, 
Scymnodon,  and  Pristiophorus)  with  closely  related  liv- 
ing species  which  are  predominantly  benthic  and  deep 
water  forms.  Several  taxa  are  broad  ranging  bathy met- 
rically (Notorhvnchus  and  Pristiophorus)  but  collectively 
the  assemblage  is  taxonomically  right  for  deep  water.  A 
precise  depth  would  be  difficult  to  substantiate  but  all 
forms  would  be  expected  to  occur  together  at  a depth  of 
600  to  1500  ft  [180  to  460  m]. 

Silicified  otoliths  are  usually  poorly  preserved,  how- 
ever, the  majority  (98%)  are  of  mesopelagic  lantemfishes 
(Family  Myctophidae).  Many  compare  favorably  to  the 
genus  Diaphus.  The  extant  D.  theta  occurs  today  in  the 
N.  Pacific  from  N.  Baja  California  to  the  Gulf  of  Alaska 
and  Japan,  at  depths  from  surface  (over  deep  water)  to 
2600  ft  [790  m]. 

If  one  considers  only  the  present  day  bathymetric  dis- 
tribution of  the  genera  of  sharks  and  bony  fishes  known 
to  occur  at  Knappton,  the  assemblage  from  all  four  lo- 
calities would  have  to  be  characterized  as  a mix  of  epi- 
pelagic  and  deep  water  benthic  sharks  and  mesopelagic 
teleosts.  A bottom  depth  of  600  to  1500  ft  [180  to  460 
m]  would  not  be  unreasonable. 

Although  the  sample  is  small,  the  absence  of  other 
selachian  taxa  (e.g.  Heterodontus,  Squatina,  Squalus, 
Triakis,  Mustelus,  Galeorhinus,  and  assorted  skates  and 
rays)  strongly  dictates  against  both  a shallow  water  (shelf) 
origin  of  the  fauna  or  resedimentation  of  a shallow  water 
assemblage  into  deeper  water  by  turbidites  or  related 
processes. 

From  LAM  Vertebrate  Locality  4510  (=LAM  Invertebrate 
Locality  5842)  Welton  has  identified  Scymnodon  sp.,  cf.  Dia- 
phus sp.,  and  Myctophidae  with  a combined  modem  bathy- 
metric distribution  of  300  to  800  m. 

Because  Aturia  is  20  times  as  abundant  as  any  other  mol- 
lusk  in  the  fauna,  particular  attention  should  be  paid  to  its 
inferred  ecologic  requirements.  Nautilus,  the  structurally 
similar  closest  living  relative  of  Aturia,  implodes  at  a water 
depth  of  785  m (Kanie  et  al.,  1980),  which  presumably  sets 
a maximum  living  depth  for  Aturia  and  the  associated  mega- 
fauna and  microfauna.  Nautilus  eggs  are  probably  laid  in 
shallow  water,  100  m or  less.  After  hatching,  the  young  de- 
scend to  250  to  350  m (Hamada,  Obata,  and  Okutani,  1980: 
47).  The  first  seven  septa  in  Nautilus  have  shown  low  5I80 
values,  indicating  Nautilus  hatched  in  warm,  shallow  water, 
about  24°C  (Cochram,  Rye,  and  Landman,  1981:477).  The 
eighth  and  later  septa  have  higher  <5lftO  values,  suggesting 
that  the  juvenile  Nautilus  subsequently  migrates  to  deeper, 
colder  water,  about  16°C.  Indirect  evidence  for  the  need  of 
shallow  warm  water  for  egg  laying  and  hatching  of  Aturia 
lies  in  the  present  distribution  of  Nautilus  in  the  south  Pacific 


Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks  7 


Other  authors: 

Musashia  ( Musashia ) n.  sp.  Armentrout  in  MS  (1973) 

Musashia  ( Musashia ) sp.  of  Allison  and  Marincovich  (1981,  pi.  3, 
figs.  8,  14) 

Musashia  (Musashia)  sp.  of  Allison  and  Marincovich  (1981,  pi.  3, 
figs.  12,  13) 

Musashia  (Musashia)  sp.  of  Allison  and  Marincovich  (1981,  pi.  3, 
figs.  16,  17) 

Miopleiona  sp.  A Durham  (1944:178;  UCMP  35421,  35422) 
Miopleiona  sp.  B Durham  (1944:178;  UCMP  35423) 

Miopleiona  weaveri  Tegland  (1933:127-128,  pi.  11,  figs.  1-5) 
Miopleiona  scowensis  Durham  (1944:177-178,  pi.  17,  fig.  15) 
Psephaea  (Miopleiona)  cf.  P.  (M.)  weaveri  (Tegland)  of  Addicott 
(1970,  pi.  13,  figs.  15,  19) 

Psephaea  (Miopleiona)  indurata  (Conrad)  of  Moore  (1963:43-44,  pi. 

7,  figs.  1,  2,  3-9,  11;  pi.  8,  figs.  1-4,  5) 

Miopleiona  oregonensis  Dali  (1909:35-36,  pi  18,  figs.  3,  7) 
Miopleiona  sp.  Clark  (1918,  pi.  23,  fig.  13;  UCMP  11244 
Psephaea  (Miopleiona)  weaveri  (Tegland)  of  Addicott  (1970,  pi.  13, 
fig.  17) 

Miopleiona  indurata  (Conrad)  of  Clark  (1918:185;  UCMP  12030) 
Psephaea  corrugata  Clark  (1932:831,  pi.  21,  figs.  5,  11) 

Miopleiona  sp.  Loel  and  Corey  (1932:241 ; UCMP  12136) 
Psephaea  (Miopleiona)  cf.  P.  (M.)  indurata  (Conrad)  of  Addicott 
(1970,  pi.  13,  figs.  6,  8) 

Musashia  indurata  (Conrad)  of  Addicott  (1976c,  pi.  3,  fig.  27) 
Musashia  n.  sp.  of  Addicott  (1976a,  pi.  4,  fig.  18) 

Postellaria  indurata  Conrad  (1849:727-728,  pi.  19,  fig.  12) 
Miopleiona  indurata  (Conrad)  of  Weaver  (1942:491,  pi.  94,  figs.  5, 

8,  13) 


This  report: 

Musashia  (Musashia)  n.  sp.  a 
Musashia  (Musashia?)  sp.  b 

Musashia  (Musashia?)  n.  sp.  c 

Musashia  (Nipponomelon?)  n.  sp.? 

Musashia  (Musashia)  n.  sp.  a 
Musashia  sp. 

Musashia  (Nipponomelon)  weaveri  (Tegland) 

Musashia  (Nipponomelon)  weaveri  (Tegland) 

Musashia  (Nipponomelon)  weaveri  (Tegland) 

Musashia  ( Miopleiona)  indurata  (Conrad) 

Musashia  (Nipponomelon)  oregonensis  (Dali) 

Musashia  (Nipponomelon?)  sp. 

Musashia  (Nipponomelon)  weaveri  (Tegland) 

Musashia  (Nipponomelon?)  sp.  cf.  M.  (N.)  weaveri  (Tegland) 
Musashia  (Neopsephaea)  corrugata  (Clark) 

Musashia  (Nipponomelon)  shik amain,  sp. 

Musashia  (Nipponomelon)  shikamai  n.  sp. 

Musashia  (Nipponomelon)  shikamai  n.  sp. 

Musashia  (Nipponomelon)  n.  sp.? 

Musashia  (Miopleiona)  indurata  (Conrad) 

Musashia  (Miopleiona)  indurata  (Conrad) 


Figure  9.  Allocation  of  Eastern  Pacific  Tertiary  volutids  assigned  to  the  genus  Musashia. 


and  in  the  fact  that  Aturia  became  extinct  in  the  eastern 
Pacific  at  the  close  of  the  early  Miocene.  A combination  of 
cooling  and  marine  regression  may  have  eliminated  suitable 
sites  for  reproduction.  Aturia  also  probably  inhabited  a shelf 
or  slope  environment  where  it  could  have  come  into  shallow 
warm  water  for  nocturnal  feeding  and  reproduction  and  eas- 
ily returned  to  deeper  water  for  resting  and  escape  from  pred- 
ators. 

A depth  between  100  and  350  m for  the  organisms  pre- 
served in  the  concretions  is  indicated  on  the  basis  of  all  the 
mollusks  including  the  most  abundant  element  of  the  fauna, 
Aturia.  Foraminifers  from  nearby  sediment  at  the  modem 
landslide  at  Knappton  indicate  a greater  depth  (about  1000 
m).  If  the  difference  is  real,  a possible  explanation  is  that  an 
early  Miocene  submarine  landslide  transported  the  Aturia- 
bearing  sediment  into  deeper  water  before  lithification. 

Other  fossil  remains  present  in  the  biota  but  not  yet  studied 
in  detail  include  radiolarians,  coelenterates,  echinoderms, 
bryozoans,  brachiopods,  decapods,  polychaetes,  trace  fossils, 
marine  mammals  (cetaceans),  birds,  seeds,  and  wood. 

The  pelecypods  are  mostly  infaunal  at  shallow  subbottom 
depths;  the  spantagoid  echinoids  and  marine  worms  also  are 
infaunal,  perhaps  to  depths  as  much  as  6 cm.  The  gastropods 
are  mostly  epifaunal,  although  the  cassids  may  have  plowed 
through  the  sediment  in  search  of  their  echinoid  prey.  The 
sponges,  coral,  and  crabs  were  also  mostly  epifaunal. 

Temperature  data  obtained  from  living  or  closely  related 


molluscan  species  are  somewhat  ambiguous.  Whereas  mol- 
lusks such  as  Ancistrolepis,  Aforia,  Portlandia,  Acharax, 
Acesta  ( Plicacesta ),  Acesta  (Acesta),  and  Lucinoma  suggest 
temperatures  between  5 and  8°C,  Aturia  may  have  required 
a temperature  of  at  least  1 6°C. 

MOLLUSCAN  PALEONTOLOGY 

The  molluscan  fauna  consists  of  33  taxa  and  many  of  these 
have  been  treated  by  Tegland  (1931,  1933),  Durham  (1944), 
and  Addicott  (1970,  1976b,  1976c).  Taxonomic  notes  rather 
than  formal  systematic  descriptions  are  used  for  all  but  new 
species  to  avoid  redundancy.  Newly  described  species  are 
treated  more  formally. 

The  following  taxa  are  included,  and,  unless  otherwise  in- 
dicated, all  are  from  LAM  Locality  5842  (Unit  IV)  and  are 
illustrated  at  natural  size. 

Gastropods: 

“ Bathybembix"  hickmanae  n.  sp. 

Epitonium  ( Nitidiscala ?)  sp. 

Sthenorytis  sp. 

Unidentified  naticids 
Liracassis  durhami  Kanno 
Liracassis  apt  a (Tegland) 

Buccinid? 

Bruclarkia  yaquinana  (Anderson  and  Martin) 
Ancistrolepis  jimgoederti  n.  sp. 


8 Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks 


Priscofususl  sp.  cf.  P.  geniculus  (Conrad) 

Musashia  (Musashia)  n.  sp. 

Musashia  ( Nipponomelon ) shikamai  n.  sp. 

Musashia  (Miopleiona)  n.  sp. 

Aforia  wardi  (Tegland) 

Turriculal  sp. 

Megasurculal  sp.  cf.  M.  wynoocheensis  (Weaver) 
Microglyphus  n.  sp.? 

Pelecypods: 

Acila  ( Acila ) gettysburgensis  (Reagan) 

Portlandia  ( Portlandia ) chehalisensis  (Arnold) 

Acharax  dalli  (Clark) 

Limopsis  nitens  (Conrad) 

Modiolus  addicotii  n.  sp. 

Acesta  ( Acesta ) twinensis  (Durham) 

Acesta  ( Plicacesta ) wilsoni  n.  sp. 

CrassostreaP.  sp. 

Lucinoma  hannibali  (Clark) 

Thyasira  (Conchocele)  disjuncta  (Gabb) 

Nemocardiuml  sp.  cf.  N.  lorenzanum  (Arnold) 

Macoma  sp.  cf.  M.  twinensis  Clark 
Cochlodesma  bainbridgensis  Clark 
Teredinid 
Scaphopod: 

Dentalium  ( Fissidentaliuml ) sp.  cf  D.  porterensis  Weaver 
Cephalopods: 

Aturia  angustatata  (Conrad) 

Sepiid? 

ABBREVIATIONS 

CAS:  California  Academy  of  Sciences,  San  Francisco. 
LACMIP:  Natural  History  Museum  of  Los  Angeles  County, 
Invertebrate  Paleontology  Section,  California. 

LACMP:  Natural  History  Museum  of  Los  Angeles  County, 
Invertebrate  Paleontology  Section,  California. 

LAM:  Natural  History  Museum  of  Los  Angeles  County,  Cal- 
ifornia. 

CAS/SU:  Stanford  University,  Stanford,  California.  (The 
Stanford  University  collections  are  now  housed  at  the  Cal- 
ifornia Academy  of  Sciences.) 

SU:  Stanford  University,  Stanford,  California. 

UC:  University  of  California,  Berkeley. 

UCMP:  University  of  California,  Museum  of  Paleontology, 
Berkeley. 

USGS:  U.S.  Geological  Survey,  Washington,  D.C.,  Cenozoic 
locality  register. 

USGS  M:  U.S.  Geological  Survey,  Menlo  Park,  California, 
Cenozoic  locality  register. 

USNM:  National  Museum  of  Natural  History,  Washington, 
DC. 

UW:  University  of  Washington,  Seattle,  Washington. 

GASTROPODS 

Trochidae 

The  genus  Bathybembix  is  used  here  in  a broad  sense  fol- 
lowing Hickman  (1980:16)  who  is  currently  undertaking  a 


detailed  revision  of  the  large  tuberculate  trochid  gastropods 
allied  to  Bathybembix.  The  Pacific  Northwest  fossil  species, 
Turcicula  Columbiana  Dali  (1909:99-100,  pi.  3,  figs.  2,  1 1) 
and  T.  washingtoniana  Dali  (1909:99-100,  pi.  17,  figs.  1,  2; 
pi.  18,  fig.  4)  were  considered  by  Rehder  (1955:225)  to  “be- 
long to  Bathybembix,  or  are  more  closely  related  to  that  genus 
than  to  any  other.”  Noda  ( 1 975:60)  believed  that  “ Turcicula ” 
Columbiana  and  “7’.”  washingtoniana  differ  sufficiently  to 
warrant  a new  subgeneric  name.  Certainly  “77”  washingtoni- 
ana needs  more  careful  scrutiny  in  terms  of  generic  or  subge- 
neric allocation.  Other  Pacific  coast  fossil  species  that  have 
been  assigned  to  Turcicula  or  to  Bathybembix  are:  Turcicula 
arnoldi  Durham  (1944:153-154,  pi.  15,  fig.  10),  Turcicula 
sanctacruzana  Arnold  (1908:373,  pi.  33,  fig.  4),  Turcicula 
turbonata  Clark  (1932:826,  pi.  20,  fig.  1 1),  and  Bathybembix 
nitor  Hickman  (1980:17-18,  pi.  2,  figs.  1,  2).  In  addition, 
Armentrout  (1973),  in  his  study  of  the  Lincoln  Creek  For- 
mation in  Washington,  recognized  three  new  species  which 
he  assigned  to  Bathybembix. 

“ Bathybembix ” hickmanae  n.  sp. 

Figures  10-12,  18 

Bathybembix  aff.  B.  arnoldi  (Durham).  Addicott,  1976b,  figs. 

6u,  w. 

“ Bathybembix ” hickmanae  is  a thin-shelled,  moderately  large 
trochid  with  five  whorls.  The  body  whorl  is  characterized  by 
two  spiral  cords  separated  by  an  almost  vertical  angulation. 
The  whorls  of  the  spire  also  have  two  spiral  cords,  one  at 
the  suture  separated  from  the  cord  above  by  the  same  type 
of  vertical  angulation.  The  outer  shell  layer  is  preserved  only 
in  small  patches,  but  the  spirals  may  have  been  keeled  and 
tuberculate  on  the  shoulder  of  the  body  whorl  and  the  spire 
whorls;  the  rest  of  the  shell  may  have  been  smooth.  The 
available  specimens  are  poorly  preserved  and  no  nacreous 
shell  material  is  apparent  on  any  of  shell  patches  preserved, 
whereas  nacreous  shell  material  is  commonly  preserved  on 
both  “ Bathybembix ” columbiana  and  “ Bathybembix ” wash- 
ingtoniana. 

HOLOTYPE.  LACMIP  6623,  height  42  mm,  width  35 
mm;  paratypes  LACMIP  6621,  height  41  mm,  width  35  mm; 
LAM  6622,  height  28  mm. 

TYPE  LOCALITY.  LAM  5842. 

“ Bathybembix ” hickmanae  somewhat  resembles  “R.”  ar- 
noldi (Durham)  and  “B.”  sanctacruzana  (Arnold)  in  outline 
but  differs  in  having  both  a steeper  and  longer  slope  between 
the  spirals  and  the  suture  on  the  body  whorl.  In  addition, 
“R.”  hickmanae  has  the  anterior  spiral  on  the  spire  whorls 
at  the  suture,  whereas  “2?.”  arnoldi  and  “B.”  sanctacruzana 
have  a space  between  the  anterior  spiral  and  the  suture.  The 
vertical  angulation  between  spirals  on  the  body  whorl  and 
the  higher  spire  in  proportion  to  width  separate  “2?”  hick- 
manae from  “5.”  washingtoniana  (Dali).  In  addition,  “2?” 
washingtoniana  has  a strong  keel  on  the  periphery  of  the 
body  whorl.  The  vertical  angulation  between  spirals  on  the 
body  whorl  distinguishes  “ B .”  hickmanae  from  “2?”  tur- 
bonata (Clark).  “ Bathybembix ” columbiana  (Dali,  1 909: 1 00, 
pi.  3,  figs.  2,  11)  has  a higher  spire  and  larger  nodes  than 


Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks  9 


“5.”  hickmanae.  “ Bathybembix”  hickmanae  differs  from 
“ Bathybembix”  nitor  (Hickman)  in  having  a quadrate  rather 
than  an  ovate  aperture. 

Traditionally,  the  species  assigned  to  Bathybembix  have 
been  assumed  to  indicate  deep,  often  bathyal  depths.  The 
bathymetric  distribution  of  living  species  in  Japan,  assigned 
to  Turcicula,  Bathybembix,  Ginebis,  and  Convexia,  is  100  to 
1000  m (Noda,  1975:58,  fig.  3). 

This  species  is  named  in  honor  of  Carole  S.  Hickman. 

Epitoniidae 

Epitonium  ( Nitidiscala ?)  sp. 

Figures  13,  14,  16 

A latex  impression  of  the  mold  of  a specimen  preserved  in 
a concretion  is  illustrated  along  with  the  original  external  and 
internal  molds.  It  is  a thick-shelled  epitoniid  with  seven  whorls 
preserved  each  of  which  probably  bore  14  or  15  thick  axial 
ribs  (seven  are  exposed).  No  spiral  sculpture  is  preserved. 

The  rounded  whorls,  deep  sutures,  slim  high  spire,  and 
lack  of  spiral  sculpture  between  the  axial  ribs  suggest  Niti- 
discala. 

This  species  somewhat  resembles  Epitonium  ( Cirsotrema ) 
saundersiEe^and  (1933:133,  pi.  13,  figs.  7-9;  Durham,  1937: 
491-492,  pi.  57,  fig.  21)  which  occurs  in  the  Liracassis  rex 
Molluscan  Zone  (Durham,  1944: 1 58).  Epitonium  saundersi, 
however,  has  spiral  sculpture  and  less  rounded  but  wider 
whorls  than  E.  (TV.?)  sp. 

Sthenorytis  sp. 

Figures  15,  19,  20 

Sthenorytis  sp.  may  have  had  five,  or  possibly  more,  rapidly 
enlarging  well-rounded  whorls,  including  a very  large  body 
whorl  set  off  from  the  axis  at  an  angle  of  about  40°,  as  is 
typical  of  Sthenorytis.  The  suture  presumably  is  deep,  and 
the  body  whorl  may  have  had  16  varices.  The  varices  are 
rather  evenly  spaced,  triangular  in  cross  section,  sharp  edged, 
and  project  about  4 mm  beyond  the  shell. 

The  only  described  species  resembling  S.  sp.  is  Sthenorytis 
ventricosum  (Clark,  1918:164,  pi.  23,  fig.  14)  from  the  San 
Ramon  Sandstone,  California.  It  has  a smaller  body  whorl 
(31  mm  wide)  than  S.  sp.  (36  mm  wide)  and  is  45  mm  high 
compared  to  about  55  mm  for  S’,  sp.  The  varices  number 


about  1 2 and  are  rounded  on  S.  ventricosum;  S.  sp.  has  about 
1 6 varices  that  are  triangular  and  sharp-edged. 

Three  Pacific  coast  Tertiary  species  are  assigned  to  Sthe- 
norytis: S.?  crescentense  (Durham,  1937),  S.  ventricosum 
(Clark,  1918),  and  S.  stearnsi  (Dali,  1 892).  The  geologic  range 
of  these  species  is  Eocene  to  Pliocene  and  the  geographic 
range  is  northwestern  Washington  to  southern  California. 

Sthenorytis  lives  today  no  farther  north  than  the  Gulf  of 
California  and  Cape  San  Lucas;  it  lives  in  warm  water  in  the 
Pacific  and  the  Atlantic  and  is  usually  found  on  sandy  bot- 
toms (Durham,  1937:499).  Woodring  (1959: 184)  reported  S. 
pernobilis  (Fischer  and  Bemardi)  from  Cape  Hatteras  to  the 
Lesser  Antilles  at  depths  of  134  to  220  m,  Keen  (1971:434, 
436)  recorded  5”.  dianae  (Hinds)  from  Baja  California  Sur  in 
82  to  145  m and  S.  turbinum  (Dali)  from  the  Gulf  of  Cali- 
fornia to  the  Galapagos  Islands  in  82  to  550  m,  and  Clench 
and  Turner  (1950:225-226)  recorded  S.  pernobilis  (Fischer 
and  Bemardi)  from  North  Carolina  to  the  Lesser  Antilles  at 
134  to  284  m. 

Naticidae 

Unidentified  naticid 

Figure  17 

Naticids  that  may  represent  Polinices  ( Euspira ) are  repre- 
sented by  three  specimens,  none  with  the  umbilical  area  well 
enough  preserved  for  positive  identifications. 

Cassididae 

Two  species  of  Liracassis,  L.  durhami  and  L.  apt  a,  were 
found  in  the  upper  part  of  the  Lincoln  Creek  Formation. 

Liracassis  durhami  Kanno 

Figures  21-23,  25,  26 

Liracassis  durhami  Kanno,  1971:112-113,  pi.  13,  figs. 
14a-b. 

Liracassis  durhami  Kanno  has  nodes  on  the  shoulder  that 
usually  form  oblique  ridges  to  the  suture  (Figs.  21-23,  25), 
but  may  be  separated  from  the  suture  by  a narrow  unsculp- 
tured band  or  confined  to  two  spiral  straps.  Secondary  spirals 
are  absent  on  the  body  whorl  except  for  one  specimen  which 


Figures  10-23.  “ Bathybembix ” hickmanae  n.  sp.,  Epitonium  ( Nitidiscala ?)  sp.,  Sthenorytis  sp.,  unidentified  naticid,  and  Liracassis  durhami 
Kanno. 

Figures  10-12,  18.  “ Bathybembix ” hickmanae  n.  sp.  10.  Paratype  LACMIP  6621,  height  41  mm,  width  35  mm.  Showing  cross  section 

of  spiral  cords.  11.  Paratype  LACMIP  6622,  height  28  mm.  Showing  nodes  on  shoulders  of  spire  and  body  whorl,  x 1.5.  12,  18.  Holotype 
LACMIP  6623,  height  42  mm,  width  35  mm.  Showing  patches  of  preserved  shell. 

Figures  13,  14,  16.  Epitonium  ( Nitidiscala ?)  sp.  13.  Latex  impression  of  specimen  shown  in  Figure  16,  xl.5.  14.  Internal  mold,  height 
25  mm,  width  10  mm.  LACMIP  6624a.  16.  External  mold  from  which  latex  impression  was  taken  shown  in  Figure  13,  x2.0;  height  26  mm, 
width  10  mm.  LACMIP  6624. 

Figures  15,  19,  20.  Sthenory’tis  sp.  15.  View  looking  down  upon  spire,  x 1.5;  height  55  mm.  LACMIP  6625a.  19.  View  of  base  of  same 
specimen  in  Figure  15  showing  triangular  cross  section  of  varices,  x 1.5.  LACMIP  6625b.  20.  Portion  of  body  whorl  of  specimen  shown  in 
Figure  19,  width  37  mm. 

Figure  17.  Unidentified  naticid,  x 1.5;  height  23  mm,  width  20  mm.  LACMIP  6626. 

Figures  21-23.  Liracassis  durhami  Kanno.  21.  Showing  one  secondary  rib  in  interspace,  x 1.5;  width  46  mm.  LACMIP  6627.  22.  Showing 
ridges  on  shoulder  of  body  whorl;  height  57  mm,  width  46  mm.  LACMIP  6628.  23.  Showing  spire;  height  63  mm,  width  48  mm.  LACMIP 
6493. 


10  Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks 


Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks  1 1 


has  a single  intercalary  in  one  interspace  (Fig.  26).  Liracassis 
durhami  always  has  a rounded  body  whorl  without  nodes 
below  the  shoulder;  this  character  distinguishes  it  from  Lira- 
cassis  rex  (Tegland,  1931:413-415,  pi.  60,  fig.  12;  pi.  61,  figs. 
1-4;  pi.  62,  figs.  1-6),  depicted  in  Figure  32  and  the  tabulate 
form  of  L.  apta  (Figs.  27,  31,  33,  39,  42).  From  the  round 
form  of  L.  apta  (Figs.  24,  28,  30,  47),  L.  durhami  is  distin- 
guished by  its  more  concave  body-whorl  shoulder,  ridgelike 
nodes  on  the  shoulder,  and  the  absence  of  intercalaries  on 
the  body  whorl.  Liracassis  durhami  always  has  nodes  on  the 
shoulder  of  the  body  whorl,  whereas  the  round  form  of  L. 
apta  may  not.  Liracassis  petrosa  (Moore,  1963,  pi.  10,  figs, 
7,17)  differs  from  L.  durhami  in  having  intercalaries  between 
primary  spirals  on  the  body  whorl  and  nodes  that  are  more 
fluted  on  the  periphery  of  the  body-whorl  shoulder,  forming 
less  oblique  and  generally  smaller  ridges  on  the  shoulder. 

Liracassis  durhami  ranges  from  the  northeastern  Gulf  of 
Alaska  to  southwestern  Washington;  it  occurs  in  the  upper 
part  of  the  Poul  Creek  Formation  and  the  lower  part  of  the 
Yakataga  Formation,  Alaska,  as  well  as  in  the  upper  part  of 
the  Lincoln  Creek  Formation,  Washington.  In  the  lower  part 
of  the  Yakataga  Formation  in  Alaska,  L.  durhami  is  asso- 
ciated with  Pillarian  mollusks  (Scott  McCoy,  pers.  commun., 
1980).  In  Washington,  L.  durhami  ranges  from  the  upper 
Galvinian  Stage  through  the  Juanian  Stage.  The  infrequent 
occurrence  of  L.  durhami  compared  to  L.  apta  in  the  upper 
part  of  the  Lincoln  Creek  Formation  may  indicate  that  south- 
western Washington  was  at  the  southern  limit  of  its  range. 
Liracassis  apta  is  restricted  to  the  Juanian,  but  has  a greater 
geographic  range  than  L.  durhami,  having  been  found  as  far 
south  as  California  (Ham,  1952:8). 

Liracassis  apta  (Tegland) 

Figures  24,  27-31,  33,  39,  42,  46,  47 

Galeodea  apta  Tegland,  1931:415-417,  pi.  63,  figs.  1-10. 

Liracassis  apta  is  common  in  the  upper  part  of  the  Lincoln 
Creek  Formation.  The  largest  specimens  of  L.  apta  usually 
have  round  shoulders  (Figs.  28,  30),  but  it  is  not  uncommon 
to  find  large  specimens  with  tabulate  shoulders  nor  is  it  un- 
usual to  find  small  specimens  with  round  shoulders.  The 
largest  specimens  of  the  round-shouldered  form  almost  never 
have  nodes  on  either  the  shoulder  or  the  rest  of  the  body 
whorl.  The  tabulate  form  always  has  nodes  on  the  shoulder 
of  the  body  whorl  and  may  have  them  on  spiral  cords  anterior 
to  the  shoulder.  Of  the  Liracassis  species  studied  (Moore, 
1984),  L.  apta  is  the  only  one  showing  the  wide  range  of 
variation  first  noted  by  Tegland  (1931:401,  406),  who  dis- 


tinguished three  varieties.  Tegland’s  varieties  1 and  2 fit  into 
the  tabulate  form  and  variety  3 into  the  round-shouldered 
form.  Liracassis  apta  has  intercalaries  in  the  interspaces  be- 
tween primary  spiral  cords  (Fig.  24)  as  do  L.  rex,  from  the 
type  section  of  the  Blakeley  Formation  of  Weaver  (1912), 
Washington,  and  L.  petrosa,  from  the  Astoria  Formation, 
Oregon,  but  L.  durhami  is  distinguished  by  having  smooth 
spaces  between  the  primary  spirals.  Liracassis  rex  always  has 
nodes  on  the  concave  shoulder  of  its  body  whorl;  these  nodes 
are  not  confined  to  the  shoulder  but  continue  adapically  as 
ridges  to  the  suture,  whereas  in  L.  apta  the  nodes,  if  present, 
are  confined  to  the  shoulder  periphery.  Liracassis  rex  always 
has  at  least  one,  and  commonly  more  than  one,  row  of  nodes 
below  the  shoulder;  L.  apta  never  has  more  than  one  row 
and  commonly  has  none  at  all.  The  nodes  on  the  periphery 
of  the  shoulder  of  L.  petrosa  are  fluted,  whereas  they  are 
rounded  in  L.  apta.  Liracassis  apta  is  restricted  to  the  Jua- 
nian Stage,  and  its  geographic  range  is  from  California  to  the 
western  Gulf  of  Alaska. 

Studies  of  living  cassids  have  shown  that  these  carnivores 
eat  only  echinoids,  and  that  different  species  within  each 
cassid  genus  prefer  certain  distantly  related  genera  of  echi- 
noids. That  different  echinoids  can  serve  as  food  for  closely 
related  cassids  makes  clear  how  two  different  species  of  Lira- 
cassis, such  as  L.  durhami  and  L.  apta,  could  live  in  the  same 
biologic  community,  but  in  different  niches. 

Buccinidae? 

Unidentified  buccinid 

Figure  36 

A single  specimen  of  a poorly  preserved  gastropod  may  be 
a buccinid.  Although  many  gastropod  families  can  be  elim- 
inated from  consideration  on  the  basis  of  outline,  sculpture, 
suture,  aperture,  or  siphonal  canal,  the  specimen  is  not  iden- 
tified with  complete  confidence  as  a buccinid.  The  evenly 
rounded  whorls,  slightly  impressed  suture,  elongate-oval  ap- 
erture, seeming  lack  of  columellar  plaits,  a probably  short, 
straight  siphonal  canal,  and  cancellate  sculpture  suggest  buc- 
cinid genera  such  as  Cymatophos  or  Antillophos. 

Neptuneidae 

Bruclarkia  yaquinana  (Anderson  and  Martin) 

Figures  41,  44 

Agasoma  yaquinana  Anderson  and  Martin,  1914:75,  pi.  4, 
figs.  5a-b. 


Figures  24-33.  Liracassis  apta  (Tegland),  Liracassis  durhami  Kanno,  and  Liracassis  rex  (Tegland). 

Figures  24,  27-31,  33.  Liracassis  apta  (Tegland).  24.  Round  form  without  body-whorl  nodes;  height  44  mm,  width  37  mm.  LACMIP 

6495.  27.  Tabulate  form;  height  55  mm.  LACMIP  6629.  28.  Round  form;  height  85  mm,  width  62  mm.  LACMIP  6630.  29.  Showing  round 
body  whorl  and  tabulate  spire;  height  63  mm,  width  49  mm.  LACMIP  6631.  30.  Round  form;  height  63  mm.  LACMIP  6632.  31.  Tabulate 
form  with  nodes  on  body  whorl;  height  47  mm,  width  42  mm.  LACMIP  6633.  33.  Tabulate  form  with  beads  on  body  whorl;  height  40  mm, 
width  33  mm.  LACMIP  6634. 

Figures  25,  26.  Liracassis  durhami  Kanno.  25.  Showing  smooth  interspaces  and  ridges  on  shoulder;  height  43  mm,  width  39  mm.  LACMIP 

6496.  26.  Showing  straplike  spiral  cords;  height  70  mm,  width  66  mm.  LACMIP  6494. 

Figure  32.  Liracassis  rex  (Tegland).  Showing  nodes  on  body  whorl  of  holotype  UCMP  32067;  height  75  mm,  width  64  mm.  Blakeley 
Formation,  Restoration  Point,  Seattle,  Washington. 


12  Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks 


32 


Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks  13 


Bruclarkia  yaquinana  is  represented  by  a single  incom- 
pletely preserved  specimen  (Figs.  41.  44).  As  noted  by  Ad- 
dicott  ( 1 970:90-9 1 ; 1 976c:23),  B.  yaquinana  is  characterized 
by  four  or  five  coarsely  noded  spiral  cords  on  the  body  whorl. 
Bruclarkia  oregonensis  (Conrad,  1848:433,  fig.  13;  Moore, 
1963:  pi.  3,  figs.  2,  3,  8.  11,  13)  has  finer  spiral  sculpture  and 
is  a larger,  more  rounded  species.  In  the  San  Joaquin  Valley, 
California,  the  stratigraphic  occurrence  of  B.  yaquinana  (bas- 
al part  of  the  Jewett  Sand)  and  B.  oregonensis  (Olcese  Sand) 
is  mutually  exclusive  (Addicott,  1970:91).  In  the  Clallam 
Formation  of  northwestern  Washington,  B.  oregonensis  oc- 
curs almost  exclusively  at  the  top  of  the  formation,  and  B. 
yaquinana  at  and  near  the  base  (Addicott,  1 976c:23-24). 

Ancistrolepis  jimgoederti  n.  sp. 

Figures  34,  35,  37,  38,  40,  43,  45,  56,  58,  59 
The  shell  of  Ancistrolepis  jimgoederti  is  large  and  thin  with 
eight  or  nine  subtabulate  whorls  that  bear  T-shaped  spiral 
cords  that  are  strongly  undercut  and,  on  the  holotype  (Figs. 
35,  37,  38),  preserve  three  secondary  spiral  cords.  The  in- 
terspaces on  the  body  whorl  bear  secondary  spiral  cords  and 
one  specimen  has  four  moderately  prominent  and  three  very 
fine  secondary  cords  preserved  in  one  interspace.  On  this 
same  specimen,  fine  vertical  striations  are  preserved  in  one 
interspace,  perhaps  reflecting  the  periostracum.  The  speci- 
men with  the  most  whorls  preserved  (eight)  is  68  mm  in 
height  (incomplete),  and  the  specimen  with  the  largest  body 
whorl  has  a maximum  width  of  36  mm.  The  largest  T-shaped 
spiral  cord  preserved  projects  3.2  mm  beyond  the  shell  body. 
The  T-shaped  spirals  are  preserved  only  in  concretionary 
matrix  (Fig.  58)  from  which  they  were  subsequently  exposed 
by  preparation  (Figs.  34,  35,  37,  38,  40).  The  body  of  the 
shell  is  so  thin  that  the  spiral  cords  exfoliate  easily  from  the 
shell,  leaving  preserved  a completely  erroneous  shell  outline 
(Figs.  43,  45).  The  T-shaped  spirals  are  now  replaced  by 
sparry  calcite,  but  do  not  seem  to  have  been  hollow  folds  of 
the  shell,  as  in  the  genus  Ecphora  from  the  Miocene  of  the 
eastern  United  States  (Vokes,  1957,  pi.  25,  fig.  2). 

HOLOTYPE.  LACMIP  6636,  height  55  mm,  width  35 
mm;  paratypes  LACMIP  6635,  height  40  mm,  width  29  mm; 
LACMIP  6637,  height  67  mm,  width  35  mm;  LACMIP  6638, 
height  65  mm,  width  35  mm;  LACMIP  6646,  width  35  mm; 
and  LACMIP  6647,  height  70  mm. 

TYPE  LOCALITY.  LAM  5842. 

Species,  such  as  Ancistrolepis  clarki  (Tegland,  1933:131  — 


132,  pi.  12,  fig.  14),  A.  landesi  (Tegland,  1933:132-133,  pi. 

13,  figs.  1-4),  and  A.  clarki  teg/andae  (Durham,  1944:177, 
pi.  17,  fig.  2),  may  also  have  had  T-shaped  spirals  that  were 
removed  by  exfoliation.  The  concretionary  fragments  re- 
maining with  the  holotype  of  A.  clarki  teglandae,  and  the 
specimen  itself  (Figs.  52,  54),  show  no  indication  of  T-shaped 
spirals,  but  some  specimens  of  A.  jimgoederti  also  show  no 
indication  of  these  spirals  (Fig.  43). 

Grant  and  Gale  (1931:657)  noted  the  similarity  in  spiral 
sculpture  between  Ancistrolepis  and  Beringius  and  suggested 
that  perhaps  Ancistrolepis  should  be  considered  a section  or 
synonym  of  Beringius  (Dali,  1887:304;  type  species  Chry- 
sodomus  crebicostatus  Dali).  Clifford  M.  Nelson,  U.S.  Geo- 
logical Survey,  kindly  called  my  attention  to  the  fact  that  the 
whorl  proportions,  aperture,  and  fasciole  of  A.  jimgoederti 
do  not  fit  Neptunea  in  the  strict  sense,  and  that  the  species 
is  more  closely  related  to  Ancistrolepis.  Species  from  the 
eastern  Pacific  Tertiary  that  have  been  assigned  to  Ancistro- 
lepis are:  Ancistrolepis  rearensis  (Clark,  1932),  Ancistrolepis 
macneili  Kanno  (1971),  Ancistrolepis  clarki  clarki  Tegland 
(1933),  Ancistrolepis  clarki  teglandae  Durham  (1944),  An- 
cistrolepis landesi  Tegland  ( 1 933),  and  Ancistrolepis packardi 
Durham  (1944). 

The  spiral  ribs  on  the  penultimate  whorl  of  Ancistrolepis 
rearensis  (C lark,  1932:831,  pi.  20,  figs.  14,  1 5)  are  described 
as  having  a fairly  prominent  collar  and  thus  would  resemble 
those  on  A.  jimgoederti.  Ancistrolepis  rearensis  has  convex 
whorls  rather  than  subtabulate  ones  and  lacks  the  concavity 
just  below  the  suture,  present  on  A.  jimgoederti.  Kanno  (1971: 
1 18)  placed  A.  clarki  teglandae  into  synonymy  with  A.  rea- 
rensis without  discussion,  and  Addicott  (1976c:23)  cited  it 
as  a junior  synonym.  Ancistrolepis  clarki  teglandae  (Figs.  52, 
54)  has  a round  body  whorl  and  a shorter,  wider  spire  than 
A.  jimgoederti.  Ancistrolepis  macneili  Kanno  (1971:1  19,  pi. 

14,  fig.  7)  is  much  more  inflated  and  has  a shorter  spire  in 
proportion  to  the  body  whorl  than  does  A.  jimgoederti.  The 
body  whorl  of  A.  clarki  clarki  is  evenly  rounded  to  the  suture; 
the  body  whorl  of  A.  jimgoederti  is  deeply  concave  between 
the  suture  and  the  first  spiral  cord  and  the  whorls  of  the  spire 
are  more  tabulate.  Ancistrolepis  clarki  clarki  is  also  wider 
and  has  fewer  whorls  (six  or  seven)  than  A.  jimgoederti.  An- 
cistrolepis landesi  has  a high  spire,  similar  to  A.  jimgoederti, 
but  the  body  whorl  is  convex  to  the  suture  rather  than  con- 
cave just  below  the  suture.  In  addition,  the  primary  spiral 
cords  on  the  body  whorl  of  A.  landesi  are  grouped  together 


Figures  34  47.  Ancistrolepis  jimgoederti  n.  sp.,  unidentified  buccinid,  Liracassis  apta  (Tegland),  and  Bruclarkia  yaquinana  (Anderson  and 
Martin). 

Figures  34,  35,  37,  38,  40,  43,  45.  Ancistrolepis  jimgoederti  n.  sp.  34.  Paratype  LACMIP  6635.  Showing  secondary  spirals,  x 1.5;  height 
40  mm,  width  29  mm.  35,  37,  38.  Holotype  LACMIP  6636;  height  55  mm,  width  35  mm.  Figures  35  and  37  show  primary  spirals  sculptured 
by  secondaries.  Figure  38  is  the  same  specimen  tipped  to  display  undercut  spiral  cords.  40.  Same  specimen  as  Figure  34.  Showing  angular 
spiral  cord,  x 1.5.  43.  Paratype  LACMIP  6637.  Showing  cast  from  which  shell  has  exfoliated;  height  67  mm,  width  35  mm.  45.  Showing  spiral 
cords  in  concretion;  height  65  mm,  width  35  mm.  Paratype  LACMIP  6638. 

Figure  36.  Unidentified  buccinid,  x2.0;  height  28  mm,  width  13  mm.  USGS  Loc.  M 7891,  USNM  363986. 

Figures  39,  42,  46,  47.  Liracassis  apta  (Tegland).  39.  Tabulate  form  with  body  whorl  nodes;  height  45  mm,  width  35  mm.  LACMIP  6497. 
42.  Tabulate  form  showing  nodes  and  intercalaries;  height  49  mm.  LACMIP  6639.  46,  47.  Rear  and  apertural  view  of  round  form;  height  68 
mm,  width  45  mm.  LACMIP  6640. 

Figures  41,  44.  Bruclarkia  yaquinana  (Anderson  and  Martin).  Views  of  aperture  and  side  showing  configuration  and  sculpture,  xl.5; 
height  30  mm,  width  22  mm.  LACMIP  6641. 


14  Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks 


on  the  middle  of  the  body  whorl  whereas  they  are  equally 
spaced  from  the  suture  on  A.  jimgoederti.  The  entire  shell  of 
A.  landesi  bears  fine,  evenly  spaced  secondary  spiral  sculp- 
ture. The  one  specimen  of  A.  jimgoederti  that  has  the  sec- 
ondary spiral  sculpture  preserved  (Figs.  34,  37,  40)  shows 
the  spirals  to  be  of  unequal  strength  and  spacing.  The  living 
species  Ancistrolepis  grammatus  (Dali,  1907:158;  1925:3,  pi. 
30,  fig.  8)  has  T-shaped  spiral  cords  on  a thin  shell  with 
subtabulate  whorls.  The  body  whorl,  however,  has  nine 
T-shaped  spirals  of  almost  equal  width  compared  to  five  on 
A.  jimgoederti  and  the  spire  whorls  have  five  rather  than 
three  or  four  spirals.  The  siphonal  canal  of  A.  jimgoederti  is 
probably  slightly  longer  and  more  strongly  recurved  than  on 
A.  grammatus  and  also  bears  finer  spiral  sculpture. 

Weaver  (1942:427)  assigned  Ancistrolepis  clarki  clarki  to 
Neptunea,  and  renamed  it  N.  teglandae,  as  N.  clarki  was  a 
homonym  of  Neptunea  clarki  (Meek).  The  reassignment  of 
A.  clarki  clarki  to  Ancistrolepis  makes  this  action  by  Weaver 
no  longer  necessary. 

Ancistrolepis  landesi  and  A.  clarki  clarki  occur  in  the  type 
Blakeley  Formation  in  the  Liracassis  rex  Molluscan  Zone. 
Ancistrolepis  clarki  teglandae  occurs  in  the  upper  part  of  the 
Pysht  Formation  of  the  Twin  River  Group  in  the  Liracassis 
apta  Molluscan  Zone.  Ancistrolepis  rearensis  was  originally 
collected  from  the  upper  part  of  the  Poul  Creek  Formation 
in  the  Liracassis  apta  Molluscan  Zone  and  the  geographic 
and  stratigraphic  ranges  were  subsequently  extended  by  Ad- 
dicott  (1976c:23)  to  the  Clallam  Formation,  Washington,  in 
the  Vertipecten  fucanus  Molluscan  Zone.  Ancistrolepis  jim- 
goederti was  collected  from  the  upper  part  of  the  Lincoln 
Creek  Formation,  in  the  upper  (Saucesian)  part  of  the  Lira- 
cassis apta  Molluscan  Zone. 

Chrysodomus  eucosimium  Dali  (1891:187-188),  the  ge- 
notype of  Ancistrolepis,  was  collected  off  the  coast  of  Un- 
alaska  in  the  Bering  Sea.  Ancistrolepis  grammatus  was  col- 
lected from  Tsugaru  Strait,  Japan,  at  a depth  of  550  m where 
the  surface  temperature  averages  about  1 8°C. 

This  species  is  named  in  honor  of  James  L.  Goedert. 

Fusinidae 

Priscofususl  sp.  cf.  P.  geniculus  (Conrad) 

Figures  50,  53,  55,  60,  61 

Priscofususl  sp.  cf.  P.  geniculus  (Conrad)  is  represented  by 
three  specimens;  two  free  of  matrix  (Figs.  50,  53,  55,  61)  and 


the  third  preserved  in  a concretion  with  the  shell  missing  on 
most  of  the  specimen,  but  replaced  by  sparry  calcite  where 
still  embedded  (Fig.  60).  Priscofusus  geniculus  (Conrad,  1849: 
728,  pi.  20,  fig.  3)  has  been  described  and  illustrated  by  Moore 
(1963:40-41,  pi.  6,  figs.  13,  15-18)  and  by  Addicott  (1970: 
101-102,  pi.  12,  figs.  21,  22,  26,  28-30)  and  occurs  in  the 
Astoria  Formation,  Oregon,  and  the  Jewett  Sand,  California. 
The  Priscofusus  reported  from  the  Clallam  Formation  (Ad- 
dicott, 1 976c:24,  pi.  2,  fig.  1 2)  and  the  Nye  Mudstone  (Moore, 
1963:41,  pi.  6,  figs.  12,  19)  may  represent  a new  species. 

Volutidae 

Neogene  volutids  of  the  eastern  Pacific  Tertiary  have  in  re- 
cent years  commonly  been  assigned  to  the  genus  Musashia 
(Hayashi,  1960)  and  the  subgenus  Musashia  or  Miopleiona 
(Dali,  1907).  A new  species  of  Miopleiona  from  the  Eugene 
Formation  in  Oregon  (Howe,  1 922)  extends  the  geologic  range 
of  that  subgenus  into  the  late  Eocene  or  early  Oligocene. 

Nipponomelon  (Shikama,  1967),  a subgenus  previously  re- 
ported from  the  Miocene  to  Holocene  in  Japan,  is  used  here 
for  most  of  the  northeastern  Pacific  volutids,  thus  extending 
the  geographic  range  of  the  subgenus  across  the  Pacific  and 
the  geologic  range  into  the  Oligocene. 

Musashia  ( Musashia ) has  a smooth  shell  or  only  thin  axial 
ribs;  axial  ribs,  if  present,  may  be  only  on  the  posterior  por- 
tion of  the  whorls  (Fig.  70);  the  suture  is  slightly  impressed. 
Musashia  (Nipponomelon)  has  thin  axial  ribs,  a slightly  im- 
pressed suture,  and  only  rarely  a sutural  collar  (Figs.  66,  68). 
Musashia  ( Miopleiona ) has  very  thick  keel-like  ribs  markedly 
curved  near  the  suture,  which  is  deeply  impressed  and  chan- 
neled (Figs.  51,  57,  64,  67).  Shikama  (1967)  thought  that 
Miopleiona  was  intermediate  between  the  subgenera  Mu- 
sashia and  Nipponomelon  and  was  uncertain  as  to  its  proper 
assignment,  although  he  placed  it  in  the  genus  Musashia.  On 
the  basis  of  the  suture  and  the  thick  axial  ribs,  Miopleiona 
could  perhaps  be  elevated  to  generic  rank.  The  type  species 
of  Miopleiona  is  Musashia  ( Miopleiona ) indurata  (Conrad, 
1849).  The  markedly  curved  axial  ribs  and  deeply  channeled 
suture  of  M.  (M.)  indurata  set  it  apart  from  all  other  described 
species  of  volutids,  but  the  undescribed  new  species  from  the 
Eugene  Formation,  Oregon  (Howe,  1922),  also  has  a deeply 
channeled  suture  (Fig.  48)  and  is  here  assigned  to  Miopleiona. 
The  suture  is  so  deeply  channeled  on  Miopleiona  that  even 
internal  molds  can  be  identified  as  belonging  in  the  subgenus. 


Figures  48-61.  Musashia  (Miopleiona)  n.  sp.,  Musashia  (Nipponomelon)  shikamai  n.  sp.,  Priscofususl  sp.  cf.  P.  geniculus  (Conrad),  Musashia 
(Nipponomelon)  indurata  (Conrad),  Ancistrolepis  clarki  teglandae  Durham,  and  Ancistrolepis  jimgoederti  n.  sp. 

Figure  48.  Musashia  ( Miopleiona ) n.  sp.  Plaster  cast  of  a specimen  from  the  Eugene  Formation,  Oregon;  height  95  mm.  CAS/SU  2358. 

Figure  49.  Musashia  (Nipponomelon)  shikamai  n.  sp.  Paratype.  Latex  impression  showing  fine  spiral  sculpture,  height  82  mm.  LACMIP 
6642. 

Figures  50,  53,  55,  60,  61.  Priscofususl  sp.  cf.  P.  geniculus  (Conrad).  50,  53.  Internal  mold  showing  traces  of  axial  ribs,  x 1.5;  height  21 
mm,  width  18  mm.  LACMIP  6643.  55,  61.  Almost  complete  internal  mold  showing  spire  outline  and  axial  ribs;  height  26  mm,  width  15 
mm.  LACMIP  6644.  60.  Showing  siphonal  canal;  height  51  mm,  width  18  mm.  LACMP  6645. 

Figures  51,  57.  Musashia  (Nipponomelon)  indurata  (Conrad).  Showing  twisted  axial  ribs  and  deeply  impressed  suture,  x0.8;  height  125 
mm,  width  59  mm.  From  the  Astoria  Formation,  Lincoln  County,  Oregon.  CAS  037058. 

Figures  52,  54.  Ancistrolepis  clarki  teglandae  Durham.  Latex  impression  of  holotype  external  molds,  UCMP  35417,  showing  spiral 
sculpture;  height  48  mm. 

Figures  56,  58,  59.  Ancistrolepis  jimgoederti  n.  sp.  56,  59.  Paratype  LACMIP  6646.  Showing  siphonal  canal,  xl.5;  width  35  mm.  58. 
Paratype  LACMP  6647.  Showing  T-shaped  cross  section  of  spiral  ribs  preserved  in  concretion,  x2.0;  height  70  mm. 


16  Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks 


58 


59 


Armentrout  (1973)  reported  an  undescribed  species  of  vo- 
lutid  from  the  Lincoln  Creek  Formation  and  assigned  it  to 
Musashia  (Musashia),  an  assignment  with  which  I concur. 
Although  several  species  of  Musashia  ( Musashia ) have  been 
described  from  the  Cenozoic  of  Japan,  Armentrout’s  material 
documents  the  occurrence  of  the  subgenus  in  western  North 
America.  It  has  a smooth  shell,  a slightly  impressed  suture 
that  is  markedly  inclined,  and  a spire  that  is  short  in  relation 
to  the  length  of  the  body  whorl  (Figs.  62,  65,  76,  80,  87). 

Subgeneric  characters  used  by  Shikama  (1967)  to  differ- 
entiate volutids,  such  as  the  character  of  the  protoconch  and 
the  number  of  initial  and  last  columellar  plaits,  are  useful 
for  the  allocation  of  living  species,  but  are  difficult  to  use 
with  incompletely  preserved  fossils.  The  character  of  the  su- 
ture and  axial  ribs  serve  best  to  distinguish  fossil  forms  both 
subgenerically  and  specifically.  On  the  basis  of  thin  axial  ribs 
and  a slightly  impressed  suture,  I assign  most  of  the  eastern 
Pacific  Tertiary  volutids  to  the  subgenus  Nipponomelon  (Fig. 
68).  My  allocation  of  eastern  Pacific  Tertiary  species  is  shown 
in  Figure  9. 

Hayashi  (1960:2)  in  his  description  of  the  genus  Musashia 
noted  that  sexual  dimorphism  is  very  pronounced  and  that 
the  large  convex  shells  may  be  female.  It  is  well  to  bear  this 
in  mind  when  looking  at  closely  related  eastern  Pacific  Ter- 
tiary species.  In  the  Knappton  fauna,  however,  slim  forms 
are  more  common  that  convex  forms,  which  would  lead  to 
the  presumably  false  conclusion  that  males  were  more  abun- 
dant than  females. 

The  oldest  recorded  occurrence  I have  found  for  Musashia 
is  Musashia  ( Nipponomelon ?)  caucasica  (Korobkov,  1949: 
694-695,  text  figs.  1,2;  1 955:205-206,  pi.  4,  figs.  6,  6a)  from 
the  middle  Eocene  in  the  Caucasus  of  the  U.S.S.R.  The  oldest 
record  of  Miopleiona  is  Musashia  (Miop/eiona)  n.  sp.  from 
the  Eugene  Formation,  Oregon,  of  late  Eocene  to  middle 
Oligocene  age.  The  oldest  record  of  Nipponomelon  in  the 
eastern  Pacific  is  in  the  lower  Oligocene  part  of  the  Lincoln 
Creek  Formation,  Washington.  Musashia  and  Nipponome- 
lon may  have  originated  in  the  western  Pacific;  Miopleiona 
is  indigenous  to  Alaska,  Washington,  Oregon,  and  California, 
and  did  not  invade  the  western  Pacific. 

Musashia  ( Musashia ) n.  sp. 

Figures  62,  65,  76,  80,  87 

Miopleiona  sp.  A Durham,  1944:178. 

Musashia  ( Musashia ) evelynae  Armentrout,  1973,  in  MS: 

338-339,  pi.  5,  figs.  25,  27. 


Musashia  ( Musashia ) n.  sp.  has  a slim  shell  with  a very  low 
spire  compared  to  the  body-whorl  height.  The  suture  is  very 
slightly  impressed  and  markedly  inclined.  The  shell  is  smooth, 
without  axial  ribs,  and  only  growth  lines  are  preserved.  The 
aperture  is  elongate  oval,  the  siphonal  fasciole  probably 
straight  and  with  a rather  thick  posterior  callus  and  with  two 
columellar  plaits,  the  anteriormost  one  bladelike. 

Musashia  ( Musashia ) n.  sp.  is  the  only  species  assigned  to 
this  subenus  in  the  eastern  Pacific.  The  type  species  of  Mu- 
sashia is  M.  ( M .)  hirasei  (Sowerby)  (Figs.  69,  70).  The  only 
other  described  species  at  all  similar  to  M.  n.  sp.  is  Musashia 
{Musashia?)  nagaoi  Shikama  (1967:1  1 1-1  12,  pi.  13,  figs.  9- 
12)  from  the  late  Oligocene  and  early  Miocene  in  Japan. 
Shikama  (1967:1  12)  considered  M.  nagaoi  to  be  unique 
among  Japanese  fulgorids  because  it  lacks  axial  ribs  and  ra- 
dial striations;  this  is  equally  true  for  M.  n.  sp.  in  the  eastern 
Pacific.  Musashia  nagaoi  has  a much  more  inflated  body 
whorl  than  M.  n.  sp.  and  the  suture  of  M.  nagaoi  is  not  as 
steeply  inclined.  Musashia  {Musashia)  n.  sp.  is  being  de- 
scribed by  J.  M.  Armentrout. 

Musashia  ( Nipponomelon ) shikamai  n.  sp. 

Figures  49,  63,  72,  74,  75,  77,  78,  82,  83,  88,  89 

Psephaea  {Miopleiona)  cf.  P.  (M.)  indurata  (Conrad).  Ad- 
dicott,  1970:105,  pi.  13,  fig.  8;  not  pi.  13,  fig.  6 {=M. 
indurata). 

Musashia  indurata  (Conrad,  1849).  Addicott,  1976c:25,  pi. 
3,  fig.  27.  Not  Rostellaria  indurata  Conrad,  1849. 

Musashia  {Nipponomelon)  shikamai  is  large,  slender,  and 
high  spired  with  about  nine  whorls.  The  shell  bears  narrow 
axial  ribs  that  are  closely  spaced  and  usually  twisted  near  the 
suture.  On  large  specimens  the  axial  ribs  disappear  toward 
the  anterior  end.  Narrow  axial  folds  between  the  ribs  pre- 
sumably represent  growth  lines;  the  entire  shell  is  sculptured 
by  closely  spaced  subrounded  spiral  cords.  The  suture  is 
slightly  impressed  and  no  subsutural  band  is  preserved.  The 
number  of  axial  ribs  ranges  from  16  to  19,  with  18  or  19 
being  the  most  common.  The  protoconch  is  not  preserved. 
The  aperture  is  assumed  to  be  elongate  oval.  The  siphonal 
fasciole  is  not  preserved  but  may  have  been  straight  and  long. 

HOLOTYPE.  LACMIP  6652,  height  135  mm,  width  52 
mm;  paratypes  LACMIP  6642,  height  87  mm;  LACMIP 
6648,  height  73  mm,  width  32  mm;  LACMIP  6649,  height 
41  mm,  width  20  mm;  LACMIP  6650,  height  90  mm,  width 
34  mm;  LACMIP  6654,  height  67  mm,  width  25  mm;  LAC- 


Figures  62-70.  Musashia  {Musashia)  n.  sp.,  Musashia  (Nipponomelon)  shikamai  n.  sp.,  Musashia  (Miopleiona)  indurata  (Conrad),  Musashia 
(Nipponomelon)  prevostiana  magna  (Kuroda  and  Habe),  and  Musashia  (Musashia)  hirasei  (Sowerby). 

Figures  62,  65.  Musashia  (Musashia)  n.  sp.  U W 16444a;  height  57  mm,  width  25  mm.  62.  Showing  smooth  shell.  65.  Showing  columnar 
plaits. 

Figure  63.  Musashia  (Nipponomelon)  shikamai  n.  sp.  Paratype  LACMIP  6648.  Showing  narrow  axial  ribs;  height  73  mm,  width  32  mm. 

Figures  64,  67.  Musashia  (Miopleiona)  indurata  (Conrad).  Showing  wide  axial  ribs  and  deeply  impressed  suture;  height  66  mm,  width  35 
mm.  USNM  363987. 

Figures  66, 68.  Musashia  (Nipponomelon)  prevostiana  magna  (Kuroda  and  Habe).  CAS  028423;  height  1 70  mm,  width  60  mm.  66.  Looking 
down  on  apex  to  show  suture.  68.  View  of  aperture. 

Figures  69,  70.  Musashia  (Musashia)  hirasei  (Sowerby).  CAS  028422;  height  165  mm,  width  56  mm.  69.  Looking  down  on  apex  to  show 
suture.  70.  View  of  aperture. 


18  Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks 


68 


69 


70 


MIP  6655,  height  104  mm,  width  36  mm;  and  LACMIP 
6651,  height  67  mm,  width  26  mm. 

TYPE  LOCALITY.  LAM  5842. 

The  species  closely  similar  to  M.  (N.)  shikamai  are  M.  ( N .) 
weaveri  (Tegland,  1933)  and  M.  (TV.)  miensis  (Araki,  1960). 
Musashia  weaveri  (Figs.  71,  81,  94)  has  a slimmer  shell  with 
fewer  (11  to  14)  and  wider  axial  ribs  and  a slightly  angulated 
shoulder  rather  than  a smoothly  rounded  one  as  in  M.  shi- 
kamai. Musashia  miensis  is  slimmer  and  more  elongate  in 
outline,  the  axial  ribs  are  not  twisted  near  the  suture,  and 
the  preserved  radial  sculpture  is  not  as  well  developed  as  in 
M.  shikamai. 

Musashia  n.  sp.  of  Addicott  (1976a:  108,  pi.  4,  fig.  18)  is 
similar  to  M.  shikamai  but  has  a few  more  axial  ribs  and  a 
concave  area  below  the  suture.  The  latter  character,  however, 
could  reflect  sexual  dimorphism. 

Musashia  ( Miopleiona ) indurata  (Conrad,  1849)  is  distin- 
guished from  M.  shikamai  by  its  deeply  impressed,  chan- 
neled suture  and  by  its  strongly  twisted  keel-like  axial  ribs. 

Musashia  indurata  (Conrad)  of  Addicott  ( 1 976c: 25,  pi.  3, 


fig.  27)  does  not  have  the  deeply  impressed,  channeled  suture 
and  wide,  keel-like  axial  ribs  of  M.  ( Mioleiona ) indurata  and 
is  here  assigned  to  M.  shikamai. 

On  the  basis  of  the  incomplete  holotype  of  Miopleiona 
scowensis  Durham  (1944:177-178,  pi.  17,  fig.  15),  I am  un- 
able to  find  any  characters  to  distinguish  it  from  M.  weaveri 
and  believe  it  should  be  synonymized  with  that  species. 

Musashia  ( Neopsephaea ) corrugata  (Clark,  1 932:831-832, 
pi.  21,  figs.  4,  5,  11;  Addicott  et  al.,  1971,  figs.  2y,  aa-bb), 
from  the  Poul  Creek  Formation,  Alaska  (Figs.  73,  90,  92, 
93,  95)  is  slimmer  than  M.  shikamai  and  has  a more  sharply 
inclined  suture  and  fewer  axial  ribs  that  are  thicker  and  more 
widely  spaced  than  in  M.  shikamai. 

Musashia  ( Musashia ) sp.  of  Allison  and  Marincovich  (1981, 
pi.  3,  figs.  12,  13;  not  pi.  3,  figs.  8,  14,  16,  17)  has  a much 
wider  body  whorl  in  proportion  to  spire  height  and  fewer 
(about  14)  axial  ribs  than  M.  shikamai.  The  specimen  figured 
by  Allison  and  Marincovich  ( 1 98 1 , pi.  3,  figs.  8,  14),  although 
poorly  preserved,  probably  belongs  in  the  subgenus  Musa- 
shia. 


Figures  71-77.  Musashia  (Nipponomelon)  weaveri  (Tegland),  Musashia  (Nipponomelon)  shikamai  n.  sp.,  Musashia  (Neopsephaea)  corrugata 
(Clark),  and  Musashia  (Musashia)  n.  sp. 

Figures  71.  Musashia  (Nipponomelon)  weaveri  (Tegland).  Showing  spacing  of  axial  ribs  and  spiral  sculpture;  height  125  mm,  width  40 
mm.  UC  locality  A 1806,  Blakeley  Formation  of  Weaver  (1912),  Bainbridge  Island,  Washington.  UCMP  35420. 

Figures  72,  74,  75,  77.  Musashia  (Nipponomelon)  shikamai  n.  sp.  72.  Showing  inflation  of  body  whorl  x0.8;  height  135  mm,  width  52 
mm.  Holotype  LACMIP  6652.  74.  Showing  spiral  sculpture  x2.0;  height  41  mm,  width  20  mm.  Paratype  LACMIP  6649.  75.  Showing 
deflection  of  ribs  near  suture;  height  90  mm,  width  34  mm.  Paratype  LACMP  6650.  77.  Showing  spacing  of  ribs  and  spiral  sculpture.  Paratype 
LACMIP  6651. 

Figure  73.  Musashia  (Neopsephaea)  corrugata  (Clark).  Showing  inclined  suture  and  sculpture;  height  120  mm,  width  33  mm.  Upper  part 
of  the  Poul  Creek  Formation,  Yakataga  Reef,  Alaska.  USNM  363988. 

Figure  76.  Musashia  (Musashia)  n.  sp.  Showing  outline  of  shell  and  inclined  suture  x 1.5;  height  52  mm,  width  20  mm.  LAM  Loc.  5843; 
LACMIP  6653. 

Figures  78-86.  Musashia  (Nipponomelon)  shikamai  n.  sp.,  Musashia ? sp.,  Musashia  (Musashia)  n.  sp.,  Musashia  (Nipponomelon)  weaveri 
(Tegland),  and  Musashia  (Nipponomelon)  prevostiana  magna  (Kuroda  and  Habe). 

Figures  78,  82,  83.  Musashia  (Nipponomelon)  shikamai  n.  sp.  78.  Paratype  LACMIP  6654.  Showing  inclined  suture  x 1.5;  height  67  mm, 
width  25  mm.  82.  Showing  narrow,  closely  spaced  axial  ribs;  height  73  mm,  width  32  mm.  Paratype  LACMIP  6648,  shown  in  Figure  63.  83. 
Paratype  LACMIP  6655.  Showing  siphonal  canal  and  spiral  sculpture;  height  104  mm,  width  36  mm. 

Figure  79.  Musashia ? sp.  Immature?  specimen  x 1.5;  height  24  mm,  width  10  mm.  LACMIP  6656. 

Figure  80.  Musashia  (Musashia)  n.  sp.  Showing  smooth  shell  and  inclined  suture;  height  57  mm,  width  22  mm.  UW  16444. 

Figures  81,  84,  86.  Musashia  (Nipponomelon)  weaveri  (Tegland).  81.  Showing  widely  spaced  axial  ribs  and  spiral  sculpture  x 1.5;  height 
40  mm,  width  23  mm.  USNM  363989.  84,  86.  From  the  basal  part  of  the  Jewett  Sand,  California.  USNM  650185;  height  73  mm,  width  30 
mm. 

Figure  85.  Musashia  (Nipponomelon)  prevostiana  magna  (Kuroda  and  Habe).  Rear  view  showing  narrow,  closely  spaced  axial  ribs  and 
suture;  height  170  mm,  width  60  mm.  CAS  028423. 

Figures  87-95.  Musashia  (Musashia)  n.  sp.,  Musashia  (Nipponomelon)  shikamai  n.  sp.,  Musashia  (Neopsephaea)  corrugata  (Clark),  and 
Musashia  (Nipponomelon)  weaveri  (Tegland). 

Figure  87.  Musashia  (Musashia)  n.  sp.  Showing  smooth  shell  and  inclined  suture  x 1.5;  height  62  mm,  width  19  mm.  USNM  363992. 
USGS  Loc.  25764,  Lincoln  Creek  Formation,  Grisdale  Quadrangle,  Washington. 

Figure  88,  89.  Musashia  (Nipponomelon)  shikamai  n.  sp.  88.  USGS  Loc.  M4050,  Clallam  Formation,  Washington.  Hypotype  (Addicott, 
1976c,  pi.  3,  fig.  27)  USNM  216000;  height  74  mm,  width  34  mm.  89.  UCMP  Loc.  3229,  Monterey  Group  of  Wagner  and  Schilling  (1923). 
Hypotype  (Addicott,  1970,  pi.  13,  fig.  6)  UCMP  12136,  a latex  impression  xl.5;  height  52  mm,  width  21  mm. 

Figures  90,  92,  93,  95.  Musashia  (Neopsephaea)  corrugata  (Clark).  90.  Paratype  UCMP  12399;  height  55  mm,  width  30  mm.  Poul  Creek 
Formation,  Alaska.  92,  95.  Holotype  UCMP  12399;  height  85  mm,  width  29  mm.  Poul  Creek  Formation,  Alaska.  93.  Showing  spire  and 
aperture  of  specimen  from  Yakataga  Reef,  Alaska;  height  120  mm,  width  33  mm.  USNM  363988.  Same  specimen  shown  in  Figure  73. 

Figures  91,  94.  Musashia  (Nipponomelon)  weaveri  (Tegland).  91.  View  looking  down  on  apex  showing  suture,  x 1.5;  height  40  mm,  width 
23  mm.  Same  specimen  shown  in  Figure  81.  USGS  Loc.  4093,  USNM  363989.  94.  Showing  outline,  axial  ribs,  and  spiral  sculpture;  height 
125  mm,  width  40  mm.  Same  specimen  shown  in  Figure  7 1 . UCMP  Locality  A 1806,  Blakeley  Formation  of  Weaver  (1912),  Bainbridge  Island, 
Washington.  UCMP  35420. 


20  Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks 


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Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks  23 


Musashia  (Musashia)  n.  sp.,  described  above,  differs  from 
M.  shikamai  by  having  a smooth  shell  without  axial  ribs  or 
radial  threads,  a much  larger  body  whorl  in  proportion  to 
height,  and  a strongly  inclined  suture. 

In  outline  and  axial  sculpture,  M.  shikamai  resembles  Mu- 
sashia {Nipponomelon)  elegantula  Shikama  (1967)  from  the 
early  Pliocene,  Japan,  but  differs  from  that  species  by  not 
having  a subsutural  band. 

The  early  whorls  of  a few  specimens  of  M.  shikamai  from 
the  upper  part  of  the  Lincoln  Creek  Formation  at  Knappton, 
Washington,  are  filled  with  barite. 

OCCURRENCE  ELSEWHERE.  Upper  part  of  the  Poul 
Creek  Formation,  Alaska,  upper  part  of  the  Pysht  Formation 
of  the  Twin  River  Group  and  the  Clallam  Formation,  north- 
western Washington,  the  Nye  Mudstone,  Oregon  (Howe, 
1922:138,  pi.  10,  fig.  3 as  Miop/eiona  clatsopensis  Howe,  n. 
sp.),  and  the  Freeman-Jewett  Silt  of  Matthews  (1955)  and 
the  Vaqueros  Formation,  southern  California. 

Musashia  ( Miopleiona ) n.  sp. 

Figure  48 

Miopleiona  n.  sp.  Howe,  1922:137. 

Musashia  ( Miopleiona ) n.  sp.,  collected  from  the  Eugene  For- 
mation in  Oregon,  is  a large  volutid  with  a thick  shell  that 
bears  perhaps  as  many  as  30  narrow,  keel-like  axial  ribs.  The 
suture  is  deeply  impressed,  channeled,  and  inclined.  The 
species  is  represented  by  a single  plaster  cast  of  a specimen 
consisting  of  half  of  two  whorls.  The  preserved  portion  is 
identical  in  all  characters  to  Musashia  ( Miopleiona ) indurata 
except  that  the  axial  ribs  are  half  as  wide  and  twice  as  closely 
spaced  as  on  M.  (M.)  n.  sp. 

There  is  no  doubt  in  my  mind  that  M.  (M.)  n.  sp.  was 
collected  from  the  Eugene  Formation  on  the  University  of 
Oregon  campus.  The  record  is  based  on  a specimen,  now 
missing  and  presumed  lost,  collected  by  Professor  Earl  L. 
Packard,  a paleontologist  of  note,  and  I have  no  reason  to 
suspect  the  locality  data.  The  specimen  was  first  mentioned 
by  Howe  ( 1 922: 1 37)  as  “ Miopleiona  n.  sp.,  (very  large),  lower 
Oligocene,  Eugene,”  and  later  by  Schenck  (1928: 1 1)  as  Mio- 
pleiona n.  sp.  from  the  Eugene  Formation,  Oregon.  A plaster 
cast  was  made  of  the  specimen  for  Schenck  and  deposited  in 
the  Stanford  University  collection,  now  housed  at  the  Cali- 
fornia Academy  of  Sciences.  Hickman  (1969)  did  not  de- 
scribe any  volutids  from  the  Eugene  Formation.  Presumably 
the  plaster  cast  was  not  seen  at  that  time,  or  the  locality 
description  was  considered  suspect. 

Musashia  ( Miopleiona ) n.  sp.  is  the  oldest  known  species 
of  Miopleiona,  as  the  subgenus  is  used  here.  Its  occurrence 
in  the  Eugene  Formation  extends  its  geologic  range  from  the 
Miocene  into  the  early  Oligocene  or  late  Eocene. 

The  Eugene  Formation  has  been  extensively  collected,  in 
part  because  it  is  exposed  in  almost  all  excavations  made  for 
buildings  on  the  University  of  Oregon  campus.  The  single, 
incomplete  specimen  of  M.  (M.)  n.  sp.  indicates  its  rareness. 
Hickman  ( 1 969:22)  suggested  that  the  Eugene  molluscan  fau- 
na lived  at  a depth  of  55  m.  The  molluscan  fauna  of  the 


upper  part  of  the  Lincoln  Creek  Formation  at  Knappton 
probably  lived  at  a depth  no  shallower  than  100  m and  the 
related  Musashia  ( Nipponomelon ) is  common  in  that  part  of 
the  unit.  Musashia  ( Miopleiona ) indurata  is  usually  found 
in  the  finer-grained,  deeper-water  facies  of  the  Astoria  For- 
mation. I suggest  that  the  molluscan  fauna  in  the  Eugene 
Formation  lived  in  shallower  water  than  was  common  for 
Miopleiona. 

Turridae 

Aforia  wardi  (Tegland) 

Figure  99 

Leucosyrinx  clallamensis  wardi  Tegland,  1933:124,  pi.  10, 
figs.  5-8. 

Aforia  wardi  is  of  medium  size  and  pagodaform,  with  nine 
strongly  angulated  whorls  and  a U-shaped  sinus  on  the  shoul- 
der (Fig.  99).  The  shell  is  smooth  above  the  angulations  but 
sculptured  by  fine  spiral  threads  below,  and  these  spirals 
extend  down  the  siphonal  canal  on  the  body  whorl.  Aforia 
campbelli  (Durham,  1944:183,  pi.  14,  fig.  4)  differs  in  having 
the  whorl  angulation  closer  to  the  suture,  and  the  angulation 
is  rounded  rather  than  bladelike.  Javidpour  (1973)  discussed 
the  phylogeny  of  eastern  Pacific  Tertiary  species  of  Aforia. 
The  correlation  diagram  (Javidpour,  1973:198,  fig.  17)  is 
misleading  in  that  Aforia  campbelli  is  shown  in  the  upper 
Oligocene  part  of  the  Lincoln  Creek  Formation,  whereas  it 
should  have  been  placed  in  the  middle  Oligocene  part  of  the 
unit  as  stated  in  the  text  (Javidpour,  1973:196,  199-200). 
Aforia  was  placed  in  the  subfamily  Turriculinae  by  McLean 
(1971:119),  following  Powell  ( 1 942). 

Living  species  of  Aforia  in  the  eastern  Pacific  have  been 
recorded  from  depths  of  6 to  2870  m (Abbott,  1 974:265)  and 
in  the  western  Pacific  from  depths  of  55  to  90  m (Kira,  1962: 
102).  Powell  (1969:411-414)  said  that  Aforia  prefers  cold 
water  ranging  from  —0.6°  to  +5.4°C  and  is  bipolar,  going 
deeper  under  equatorial  waters. 

Based  on  the  illustrations  by  Powell  (1969:411,  pi.  322, 
figs.  1-4;  414,  pi.  323,  figs.  1-3)  of  the  type  species  of  Aforia, 
Pleurotoma  circinata  Dali,  characters  such  as  apical  angle 
and  position  of  whorl  angulation  are  not  useful  in  distin- 
guishing species.  If  larger  collections  of  well-preserved  spec- 
imens of  Aforia  become  available,  future  workers  may  see 
fit  to  synonymize  some  of  the  species  proposed  for  eastern 
Pacific  Tertiary  Aforia. 

OCCURRENCE  ELSEWHERE.  Lower  part  of  the  Blak- 
eley Formation  in  the  Liracassis  rex  Molluscan  Zone,  Wash- 
ington. 

Turriculal  sp. 

Figure  97 

Turriculal  sp.  is  represented  by  one  incompletely  preserved 
specimen  on  which  the  siphonal  canal  is  not  exposed.  The 
sinus  is  U-shaped  and  confined  to  the  shoulder  slope.  The 
shell  is  sculptured  by  moderately  strong  spiral  cords  that  are 
not  noded.  Turricula  washingtonensis  { Weaver,  1912:78,  pi. 


24  Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks 


3,  fig.  31;  1942:533,  pi.  98,  figs.  16,  17,  22)  differs  from 
Turriculal  sp.  in  having  nodes. 

Megasurculal  sp. 
cf.  M.  wynoocheensis  (Weaver) 

Figure  96 

Megasurculal  sp.  cf.  M.  wynoocheensis  (Weaver)  is  repre- 
sented by  a single  poorly  preserved  and  somewhat  crushed 
specimen.  Megasurcula  wynoocheensis  (Weaver,  1912:70- 
71,  pi.  11,  figs.  87-89,  94)  is  a middle  Miocene  and  possibly 
early  Miocene  species  (Addicott,  1976c:27,pl.  3,  figs.  16,  17). 

Actenoidae 

Microglyphus ? n.  sp.? 

Figure  98 

Microglyphus ? n.  sp.?  has  three  or  possibly  four  whorls  and 
a body  whorl  that  is  very  inflated  with  the  maximum  width 
at  the  middle  of  the  whorl.  The  spiral  cords,  bounded  by 
incised  grooves,  are  not  equidimensional.  The  single  speci- 
men is  very  small  (2. 1 mm  high,  1 .9  mm  wide)  and,  although 
it  may  be  an  immature  individual,  the  number  of  whorls 
indicates  that  it  probably  is  a very  small  species.  In  addition 
to  its  small  size,  Microglyphus ? n.  sp.?  differs  from  other 
described  Tertiary  actenoids  in  having  a more  globose  body 
whorl  with  the  maximum  inflation  at  the  middle  of  the  whorl. 

PELECYPODS 

Nuculidae 

Acila  ( Acila ) gettysburgensis  (Reagan) 

Figures  100-102,  105 

Nucula  {Acila)  gettysburgensis  Reagan,  1909:171,  175,  177, 
pi.  1,  fig.  3. 

Acila  (Acila)  gettysburgensis  is  represented  by  five  specimens. 
Acila  (A.)  gettysburgensis  ranges  from  the  Matlockian  through 
the  Pillarian  Molluscan  Stages. 

Nuculanidae 

Portlandia  ( Portlandia ) chehalisensis  (Arnold) 

Figures  103,  104,  106,  107 

Malletia  chehalisensis  Arnold,  1908:365,  pi.  33,  fig.  9. 

Portlandia  ( Portlandia ) chehalisensis  is  represented  by  seven 
specimens,  some  well  preserved  (Fig.  107)  and  one  double- 
valved  (Figs.  103,  104).  One  incomplete  specimen  has  con- 
centric Saccella- like  ridges  on  the  midportion  of  the  shell 
near  the  ventral  margin  (Fig.  106).  One  single  valve  is  31.8 
mm  long,  18.5  mm  high,  and  7.0  mm  thick;  perhaps  the 
largest  specimen  of  the  species  collected.  The  largest  speci- 
men noted  by  Hickman  (1969:3 1)  measured  26  mm  in  length. 


Yoldia  reagani  Dali  (1922:306)  was  considered  a synonym 
of  Portlandia  chehalisensis  (Hickman,  1969:30). 

Living  eastern  Pacific  species  of  Portlandia  occur  no  farther 
south  than  latitude  54°N  and  are  found  at  depths  of  10  to 
2560  m and  temperatures  from  —2°  to  +6°C  (Bernard,  1983: 
13). 

Solemyidae 

Acharax  dal/i  (Clark) 

Figures  108-1 11,  114 

Solemya  dalli  Clark,  1925:73,  pi.  9,  fig.  3. 

Acharax  dalli  is  represented  by  six  specimens,  all  but  one 
double-valved.  Fingerlike  projections  of  the  periostracum  are 
partially  preserved  on  some  specimens  (Fig.  108).  Acharax 
ventricosa  (Conrad,  1849:723,  pi.  17,  figs.  7,  8),  a species 
found  in  the  Astoria  Formation  in  Oregon  and  Washington, 
is  higher  in  proportion  to  length  than  A.  dalli. 

The  eastern  Pacific  Holocene  species  Acharax  johnsoni 
(Dali,  1891)  lives  at  a depth  between  800  and  3000  m at 
temperatures  of  1°  to  9°C  (Bernard,  1983:9).  Vokes  (1955: 
536-537)  said  that  living  species  of  Acharax  are  found  at 
depths  of  5 to  3 1 80  m and  that  the  controlling  factor  in  their 
distribution  may  be  water  temperature. 

Limopsidae 

Limopsis  nitens  (Conrad) 

Figures  112,  113,  115,  116 

Pectunculus  nitens  Conrad,  1849:726,  pi.  18,  figs.  9a-b. 

Limopsis  nitens  occurs  as  numerous  single  valves  (Figs.  1 12, 
1 13)  and  occasional  paired  valves  (Figs.  115,  116).  Radial 
lines  of  sculptural  punctures  are  preserved  on  some  speci- 
mens. 

The  lithology  of  the  concretions  from  Knappton  and  the 
clustering  together  of  many  specimens  is  similar  to  the  con- 
cretion presumably  from  the  Astoria  Formation,  at  Astoria, 
Oregon,  that  contains  the  lectotype  of  L.  nitens  (Moore,  1963: 
61-62,  pi.  15,  figs.  2,  5).  Weaver  (1942:76)  suggested  that 
the  lectotype  was  collected  at  Knappton,  rather  than  at  As- 
toria, because  he  had  found  nodules  containing  large  numbers 
of  L.  nitens  at  Knappton  and  had  not  found  any  specimens 
at  Astoria.  Howe  (1922:70)  did  not  find  any  specimens  of  L. 
nitens  at  Astoria  and  I found  none  in  the  Astoria  Formation 
farther  south  (Moore,  1963:62).  The  rock  containing  the  lec- 
totype of  L.  nitens  may  have  come  from  Knappton,  or  the 
upper  part  of  the  Lincoln  Creek  Formation  may  have  been 
exposed  on  the  Columbia  River  terrace  at  Astoria  when  Dana 
made  his  collection  in  1841,  yet  no  other  mollusks  typical 
of  the  Lincoln  Creek  Formation  were  collected  by  him. 

Most  species  of  Limopsis  live  in  deep  water  (Keen,  1971: 
54);  Limopsis  diegensis  Dali  has  been  collected  at  depths  of 
120  to  1500  m and  at  temperatures  between  3°  and  27°C 
(Bernard,  1983: 1 7).  The  fact  that  Limopsis  nitens  most  com- 


Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks  25 


monly  occurs  in  clumps  suggests  that  the  species  was  gre- 
garious. 

Mytilidae 

Modiolus  addicotti  n.  sp. 

Figures  117,  125 

Modiolus  n.  sp.?  aff.  M.  restorationensis  Van  Winkle.  Ad- 
dicott,  1976c:28,  pi.  5,  fig.  5. 

Modiolus  addicotti  is  a rather  small  Modiolus  with  weakly 
inflated  valves,  a markedly  thin  shell,  and  a convex  dorsal 
margin.  The  posterior  end  is  only  moderately  enlarged,  and 
slightly  longer  near  the  ventral  margin.  The  anterior  end  is 
small  and  evenly  curved.  The  umbones  are  close  to  the  an- 
terior margin.  Patches  of  preserved  shell  are  light  brown  and 
iridescent  with  mostly  evenly  spaced  growth  lines  but  with 
a few  bunched  together  forming  low  ridges. 

HOLOTYPE.  L ACM  IP  6672,  length  47  mm,  height  25 
mm. 

TYPE  LOCALITY.  LAM  5842. 

Modiolus  addicotti  differs  from  Modiolus  restorationensis 
Van  Winkle  ( 1 9 1 8:82,  pi.  4,  fig.  5)  in  having  a convex  dorsal 
margin  and  a narrower  posterior  end. 

Modiolus  lives  intertidally  to  360  m in  the  eastern  Pacific, 
but  most  species  are  found  at  depths  no  greater  than  50  m 
(Bernard,  1983:19). 

OCCURRENCE  ELSEWHERE.  Lowermost  part  of  the 
Clallam  Formation,  northwestern  Washington. 

This  species  is  named  in  honor  of  Warren  O.  Addicott. 

Limidae 

Acesta  ( Acesta ) twinensis  (Durham) 

Figures  1 19,  123,  124 

Lima  twinensis  Durham,  1944:139,  pi.  13,  fig.  11. 

Acesta  ( Acesta ) twinensis  is  represented  by  one  incomplete 
double-valved  specimen  and  four  incomplete  single  valves 


all  of  which  are  preserved  intact  with  their  original  inflation. 
The  anterior  ears  are  small  and  well  defined  by  a deep  con- 
cave groove  along  the  anterior  margin  (Fig.  1 24).  The  anterior 
margin  is  straight,  not  concave,  and  joins  the  ventral  margin 
without  an  abrupt  break  in  alignment.  The  posterior  ears  are 
large  and  indistinctly  delineated  (Figs.  1 19,  124).  The  shells 
are  large  (maximum  estimated  height  140  mm),  thin,  and 
smooth  in  the  center  but  with  rounded  ribs  of  varying  widths 
at  the  shell  margins  (Fig.  124).  The  shells  are  inflated,  and 
the  largest  specimen  suggests  a thickness  of  25  mm  (one 
valve).  Portions  of  the  brown  translucent  outer  shell  layer 
are  preserved  on  most  specimens,  but  this  shell  layer  tends 
to  stay  attached  to  the  enclosing  rock  when  the  specimens 
are  broken  away. 

Acesta  twinensis  is  distinguished  by  its  sharply  truncated 
anterior  margin,  which  differentiates  it  from  Acesta  robertsae 
(Durham,  1944),  an  early  Oligocene  species  that  has  a more 
rounded  anterior  margin. 

Acesta  ( Acesta ) oregonensis  Clark  (1925:84,  pi.  14,  figs.  3, 
4),  a species  from  the  upper  Eocene  and  lower  Oligocene 
Keasey  Formation,  Oregon,  has  an  arcuate  anterior  margin. 

Acesta  twinensis  ranges  from  the  Matlockian  through  the 
Juanian  Molluscan  Stages.  Living  species  of  Acesta  ( Acesta ) 
are  found  in  the  eastern  Pacific  at  depths  between  600  and 
2200  m and  at  temperatures  of  1°  to  8°C  (Bernard,  1983:22). 
One  species  has  been  collected  in  the  western  Pacific  near 
Japan  at  a depth  of  185  m. 

OCCURRENCE  ELSEWHERE.  Blakeley  Formation, 
Washington. 

Acesta  ( Plicacesta ) wilsoni  n.  sp. 

Figures  118,  132,  134 

Acesta  ( Plicacesta ) wilsoni  is  of  moderate  size  and  subovate 
in  outline,  with  a thicker  shell  than  Acesta  ( Acesta ) and  radial 
ribs  of  varying  widths  and  spacing  that  are  rounded  and  most 
prominent  on  the  middle  portion  of  the  shell  but  that  persist 
to  the  shell  margins.  Beaks  small;  anterior  auricle  presumed 
small  and  delineated;  posterior  auricle  large  and  not  delin- 


Figures  96-117.  MegasurculaP  sp.  cf.  wynoocheensis  (Weaver),  Turricula ? n.  sp.?,  Microglyphus ? n.  sp.?,  Aforia  wardi  (Tegland),  Acila  ( Acila ) 
gettysburgensis  (Reagan),  Portlandia  (Portlandia)  chehalisensis  (Arnold),  Acharax  dalli  (Clark),  Limopsis  nitens  (Conrad),  and  Modiolus 
addicotti  n.  sp. 

Figure  96.  MegasurculaP  sp.  cf.  wynoocheensis  (Weaver).  Rear  view,  x 1.5;  height  32  mm,  width  20  mm.  USGS  Loc.  7891,  USNM  363990. 

Figure  97.  Turricula ? n.  sp.?  Showing  spiral  sculpture,  x2.0;  height  15  mm,  width  10  mm.  LACMIP  6657. 

Figure  98.  Microglyphus ? n.  sp.?  Showing  outline  and  spiral  sculpture,  x5.0;  height  2.7  mm,  width  2.4  mm.  LACMIP  6658. 

Figure  99.  Aforia  wardi  (Tegland).  Showing  pagodaform  outline,  x 1.5;  height  28  mm,  width  14  mm.  LACMIP  6659. 

Figures  100-102, 105.  Acila  (Acila)  gettysburgensis  (Reagan).  100.  Showing  primary  bifurcation,  x 2;  length  13  mm,  height  1 1 mm.  LACMIP 
6660.  101.  Showing  sulcus  xl.5;  height  17  mm.  LACMIP  6661.  102.  Showing  secondary  bifurcation,  x2.  LACMIP  6662.  105.  Showing 
outline  and  bifurcation,  x 1.5;  length  25  mm,  height  20  mm.  LACMIP  6663. 

Figures  103,  104,  106,  107.  Portlandia  (Portlandia)  chehalisensis  (Arnold).  103,  104.  Double-valved  specimen  showing  sculpture  and 
lunule,  x 1.5;  length  21  mm,  height  12  mm,  width  8 mm  (both  valves).  LACMIP  6664.  106.  Showing  Saccella- like  sculpture,  x2;  length  22 
mm,  height  13  mm.  LACMIP  6665.  107.  Showing  outline  and  sculpture,  x 1.5;  length  32  mm,  height  18  mm.  LACMIP  6666. 

Figures  108-111, 114.  Acharax  dalli  (Clark).  108, 114.  Showing  extensions  of  fingerlike  periostracum  and  dorsal  side;  length  28  mm,  height 
24  mm,  width  16  mm  (both  valves).  LACMIP  6667.  109,  110.  Showing  sculpture  of  double-valved  specimen,  x 1.5;  length  45  mm,  height  20 
mm,  width  1 1 mm  (both  valves).  LACMIP  6668.  111.  Showing  sculpture  x 1.5;  length  35  mm,  height  13  mm.  LACMIP  6669. 

Figures  112,  113,  115,  116.  Limopsis  nitens  (Conrad).  112.  Showing  outline,  x3;  10  mm  long,  7 mm  high.  LACMIP  6670.  113.  Showing 
radial  punctae,  x3;  length  9 mm,  height  7 mm.  LACMIP  6671.  115,  116.  Double-valved  specimen  showing  thickness  of  valves,  x5;  length 
8 mm,  height  6 mm,  width  5 mm  (both  valves).  USGS  Loc.  7891,  USNM  363991. 

Figure  117.  Modiolus  addicotti  n.  sp.  Holotype  LACMIP  6672.  Showing  outline  of  valve;  length  47  mm,  height  25  mm. 


26  Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks 


115 


110 


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Moore:  Lincoln  Creek  Formation  Moilusks  27 


eated.  No  concentric  sculpture  is  preserved.  The  holotype 
(incomplete)  is  60  mm  high  and  50  mm  wide  and  the  thick- 
ness of  one  valve  is  13  mm  (Figs.  1 18,  132).  The  paratype 
(incomplete)  is  63  mm  high  and  40  mm  wide  and  the  thick- 
ness of  one  valve  is  1 1 mm  (Fig.  134). 

HOLOTYPE.  LACMIP  6673,  length  51  mm,  height  61 
mm,  width  10  mm  (one  valve);  paratype  LAM  6686,  length 
41  mm,  height  61  mm. 

TYPE  LOCALITY.  LAM  5842. 

Acesta  wilsoni  is  smaller  than  Acesta  (Plicacesta)  oakvil- 
lensis  (Clark,  1925:84,  pi.  15,  figs.  1,  3)  and  has  fewer  ribs 
and  a more  rounded  anterior  margin.  A topotype  specimen, 
earlier  illustrated  by  Weaver  (1942,  pi.  21,  fig.  1),  is  figured 
for  comparison  (Fig.  126).  Acesta  cf.  A.  oakvi/lensis  Clark  of 
Addicott  (1976b,  figs.  6a,  c)  has  narrower  ribs  that  are  more 
widely  and  evenly  spaced  than  on  A.  wilsoni  and  may  be  a 
new  species. 

Acesta  wilsoni  and  A.  oakvi/lensis  are  the  only  Tertiary 
species  of  Plicacesta  known  in  North  America.  Acesta  n.  sp. 
of  Addicott  (1976b,  figs.  6x,  z,  ab)  may  also  be  a Plicacesta. 

The  Holocene  species  Acesta  ( Plicacesta ) sphoni  (Hertlein, 
1963)  was  collected  at  latitude  33°N  at  a depth  between  455 
and  550  m and  a temperature  between  4°  and  9°C.  Acesta 
(Plicacesta)  smithi  (Sowerby)  occurs  off  Honshu,  Japan,  at 
depths  between  90  and  185  m (Kira,  1962:145). 

This  species  is  named  in  honor  of  Edward  C.  Wilson. 

Ostreidae 

Crassostrea!  sp. 

Figure  127 

Crassostrea ? sp.  is  represented  by  a single  double-valved 
specimen  preserved  in  a concretion  and  broken  upon  re- 
moval from  the  matrix.  Indigenous  Holocene  species  of  Cras- 
sostrea live  intertidally  to  a depth  of  7 m in  the  eastern  Pacific 
(Bernard,  1983:23). 

Lucinidae 

Lucinoma  hannibali  (Clark) 

Figures  120-122 

Phacoides  ( Lucinoma ) hannibali  Clark,  1925:89,  pi.  22,  figs. 

2,  4. 

Lucinoma  hannibali  is  represented  by  six  double-valved 
specimens  (Figs.  1 20-122)  from  the  upper  part  of  the  Lincoln 


Creek  Formation,  one  with  the  shell  replaced  by  barite.  The 
specimens  range  in  height  from  27  mm  to  52  mm.  Lucinoma 
acutilineata  (Conrad,  1849:725,  pi.  18,  figs.  2,  2a,  2b)  has  a 
shorter  more  concave  dorsal  margin  than  L.  hannibali.  Vari- 
ation has  been  noted  (Moore,  1963:70)  in  the  spacing  of 
concentric  lamellae  within  single  lots  of  the  Holocene  species 
Lucinoma  annulata  (Reeve,  1850)  and  by  Addicott  (1976c: 
30)  in  the  Oligocene  to  Miocene  species  L.  acutilineata,  yet 
specimens  of  L.  hannibali  from  the  upper  part  of  the  Lincoln 
Creek  Formation  have  concentric  lamellae  rather  consis- 
tently less  densely  spaced  (Figs.  120-122)  than  the  lamellae 
on  L.  acutilineata  from  the  lower  part  of  the  Astoria  For- 
mation. Lucinoma  acutilineata  has  been  found  in  the  Eugene 
Formation  (upper  Eocene  to  middle  Oligocene)  in  Oregon 
(Hickman,  1969:38,  42)  and  in  the  lower  and  middle  Mio- 
cene Astoria  Formation  (Moore,  1963:70-71,  pi.  15,  figs.  7- 
10,  12)  in  Oregon  and  Washington.  If  L.  hannibali  and  L. 
acutilineata  are  distinct  species,  and  I believe  that  they  are, 
L.  acutilineata  may  have  preferred  somewhat  shallower  water 
(50  m or  less)  than  L.  hannibali,  and  the  two  species  coexisted 
at  different  depths.  Lucinoma  annulata  lives  today  from  lat- 
itude 3 3°  to  47°N  at  depths  of  2 5 to  750  m and  L.  aequizonata 
(Steams,  1891)  lives  from  latitude  34°  to  37°N  at  depths  of 
400  to  650  m (Bernard,  1983:29).  Lucinoma  hannibali  has 
a wider  escutcheon  and  less  concave  dorsal  margin  than  L. 
columbiana  (Clark  and  Arnold,  1923:144-145,  pi.  25,  figs. 
2a-b)  from  the  Sooke  Formation,  Vancouver  Island,  and  the 
Blakeley  Formation  of  Weaver  (1912).  Lucinoma  hannibali 
ranges  from  the  Matlockian  through  the  Juanian  Molluscan 
Stages. 

Thyasiridae 

Thyasira  ( Conchocele ) disjuncta  (Gabb) 

Figures  136,  138,  142 

Conchocele  disjuncta  Gabb,  1866:28;  1869:99,  pi.  7,  figs. 

48a-b. 

Thyasira  (Conchocele)  disjuncta  is  larger,  more  quadrate,  and 
has  a more  truncated  anterior  end  than  Thyasira  bisecta 
(Conrad,  1849:724,  pi.  17,  figs.  10,  10a)  from  the  Astoria 
Formation  in  Oregon  (Moore,  1963:72,  pi.  23,  figs.  8,  14, 
1 5).  Thyasira  disjuncta  occurs  in  the  Clallam  Formation  (Ad- 
dicott, 1976c:30,  pi.  6,  fig.  7)  and  is  living  today  (Bernard, 
1983:29).  The  presence  of  two  internal  casts  (Figs.  136,  138) 
and  one  specimen  with  the  outer  shell  preserved  (Fig.  142) 


Figures  118-125.  Acesta  (Plicacesta)  wilsoni  n.  sp.,  Acesta  ( Acesta ) twinensis  (Durham),  Lucinoma  hannibali  (Clark),  and  Modiolus  addicotti 
n.  sp. 

Figure  118.  Acesta  (Plicacesta)  wilsoni  n.  sp.  showing  outline  and  radial  ribs,  x 1.5;  length  51  mm,  height  61  mm,  width  10  mm  (one 
valve).  Holotype  LACMP  6673. 

Figures  119,  123,  124.  Acesta  (Acesta)  twinensis  (Durham).  119,  123.  Showing  posterior  ears.  119.  Length  74  mm,  height  93  mm,  width 
22  mm  (one  valve).  LACMP  6675.  123.  Length  92  mm,  height  75  mm.  LACMP  6674.  124.  Showing  anterior  ear  and  axial  ribs,  x 1.5;  length 
60  mm,  height  70  mm.  LACMP  6676. 

Figures  120-122.  Lucinoma  hannibali  (Clark).  120.  Showing  profile  of  lunule;  length  29  mm,  height  27  mm,  width  1 1 mm  (both  valves). 
LACMP  6677.  121.  Showing  sulcus;  length  39  mm,  height  34  mm,  width  16  mm  (both  valves).  LACMP  6678.  122.  Showing  concentric 
sculpture;  length  40  mm,  height  37  mm,  width  27  mm  (both  valves).  LACMP  6679. 

Figure  125.  Modiolus  addicotti  n.  sp.  Holotype  LACMP  6672.  Tipped  to  show  configuration  of  double-valved  specimen,  x 1.5;  length  47 
mm,  height  25  mm.  Same  specimen  shown  in  Figure  1 17. 


28  Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks 


in  the  upper  part  of  the  Lincoln  Creek  Formation  extends 
the  range  of  the  species  downward  into  the  upper  part  of  the 
Juanian  Molluscan  Stage.  Thyasira  disjuncta  lives  today  from 
latitude  48°  to  54°N  at  depths  of  100  to  750  m and  at  tem- 
peratures between  0°  and  7°C.  Thyasira  bisecta  lives  from 
latitude  43°  to  57°N  at  depths  between  50  and  300  m and 
temperatures  between  —1°  and  +11°C  (Bernard,  1983:29). 
The  occurrence  of  T.  disjuncta  in  the  Lincoln  Creek  For- 
mation and  of  T.  bisecta  in  the  Astoria  Formation  is  related 
to  depth  of  water;  the  Astoria  Formation  represents  a shal- 
lower-water  facies  than  the  Lincoln  Creek  Formation. 

Cardiidae 

Nemocardium ? sp.  cf.  N.  lorenzanwn  (Arnold) 

Figure  130 

Nemocardium ? sp.  cf.  N.  lorenzanum  (Arnold,  1908:366,  pi. 
33,  fig.  6)  is  represented  by  two  poorly  preserved  single  valves; 
one  is  illustrated  (Fig.  1 30). 

Tellinidae 

Macoma  sp.  cf.  M.  twinensis  Clark 

Figure  128 

Macoma  sp.  cf.  M.  twinensis  Clark  is  represented  by  a single 
valve  that  does  not  have  the  hinge  exposed  (Fig.  128).  In 


size,  outline,  and  position  of  umbo,  the  specimen  resembles 
M.  twinensis  Clark  (1925:96,  pi.  12,  fig.  7),  which  ranges 
from  late  Oligocene  to  early  Miocene. 

Periplomatidae 

Cochlodesma  bainbridgensis  Clark 

Figures  139,  141 

Cochlodesma  bainbridgensis  Clark,  1925:86,  pi.  13,  figs. 
3,  4. 

Cochlodesma  bainbridgensis  Clark  has  a thin,  internally 
nacreous,  fragile  shell,  that  is  sculptured  with  concentric  un- 
dulations (Figs.  139,  141).  Five  specimens  and  one  possible 
juvenile  (Fig.  131)  were  collected  from  the  upper  part  of  the 
Lincoln  Creek  Formation.  As  noted  by  Moore  (1976:53,  pi. 
16,  figs.  4,  6-11),  the  variation  in  outline  is  great  and  not 
useful  in  the  discrimination  of  fossil  species.  Holocene  species 
of  the  closely  related  genus  Periploma  live  intertidally  to  a 
depth  of  380  m (Bernard,  1983:64). 

Teredinidae 

Figures  172,  177,  179 

Teredinid  burrows  are  preserved  in  wood  and  the  tubes  are 
filled  with  quartz  (Fig.  1 72)  or  with  sediment  (Figs.  177,  179). 
No  pallets  are  preserved,  enabling  generic  differentiation,  but 


Figures  126-1 34.  Acesta  ( Plicacesta ) oakvillensis  (Clark),  Crassostreal  sp. , Macoma  sp.  cf.  M.  twinensis  Clark,  Flabellum  sp. , Nemocardium ? 
sp.  cf.  N.  lorenzanum  (Arnold),  Cochlodesmal  sp.,  Lima  ( Plicacesta ) wilsoni  n.  sp.,  and  Aturia  angustata  (Conrad). 

Figure  126.  Acesta  ( Plicacesta ) oakvillensis  (Clark).  Showing  closely  spaced  axial  ribs;  length  79  mm,  height  95  mm.  UC  Loc.  A368,  lower 
part  of  the  Lincoln  Creek  Formation,  Grays  Harbor  County,  Washington.  UCMP  32405. 

Figure  127.  Crassostreal  sp.  Showing  configuration;  length  70  mm,  height  102  mm.  LACMP  6680. 

Figure  128.  Macoma  sp.  cf.  M.  twinensis  Clark.  Showing  outline  and  concentric  lines,  x 1.5;  length  15  mm,  height  10  mm.  LACMP  6681. 
Figure  129.  Flabellum  sp.  x 1.5;  height  22  mm.  LACMP  6682. 

Figure  130.  Nemocardium ? sp.  cf.  N.  lorenzanum  (Arnold).  Showing  configuration  and  radial  ribs,  x 1.5;  height  20  mm.  LACMP  6683. 
Figure  131.  Cochlodesmal  sp.  Showing  outline,  x2;  length  1 1 mm,  height  8 mm.  LACMP  6684. 

Figures  132,  134.  Lima  ( Plicacesta ) wilsoni  n.  sp.  132.  Showing  outline  and  radial  ribs,  x 1.5;  length  51  mm,  height  61  mm,  width  10  mm 
(one  valve).  Holotype  LACMP  6673.  134.  Showing  radial  ribs  x 1.5;  length  41  mm,  height  61  mm.  Paratype  LACMP  6686. 

Figure  133.  Aturia  angustata  (Conrad).  Immature  specimen,  x 1.5;  height  24  mm.  LACMP  6687. 


Figures  135-145.  Aturia  angustata  (Conrad),  Thyasira  ( Conchochele ) disjuncta  (Gabb),  and  Cochlodesma  bainbridgensis  Clark. 

Figures  135,  137,  140,  143-145.  Aturia  angustata  (Conrad).  135.  Showing  outer  shell  and  growth  lines,  x0.7;  height  150  mm.  LACMP 
6688.  137.  Cross  section  showing  funnel-shaped  septal  structures;  height  29  mm.  LACMP  6689.  140.  Cross  section  showing  septal  structures; 
height  37  mm.  LACMP  6690.  143.  Apertural  view  of  broken  specimen  showing  siphuncular  orifices;  height  90  mm.  LACMP  6691.  144.  Cross 
section  showing  septal  structures;  height  60  mm.  LAM  Loc.  5843,  LACMP  6692.  145.  Side  view  showing  sutures;  height  65  mm,  width  27 
mm  (maximum  diameter).  Same  specimen  shown  in  Figure  150.  LACMP  6693. 

Figures  136,  138,  142.  Thyasira  ( Conchochele ) disjuncta  (Gabb).  136.  Showing  sulcus  and  configuration;  length  70  mm,  height  67  mm. 
LACMP  6694.  138.  Showing  sulcus  and  configuration;  length  52  mm,  height  43  mm.  LACMP  6695.  142.  Showing  outer  shell  and  concentric 
lines,  x 1.5;  length  37  mm,  height  32  mm.  LACMP  6696. 

Figures  139,  141.  Cochlodesma  bainbridgensis  Clark.  139.  Showing  concentric  undulations,  x 1.5;  length  34  mm,  height  27  mm.  LACMP 
6697.  141.  Showing  outline,  x 1.5;  length  30  mm,  height  25  mm.  LACMP  6698. 


Figures  146-154.  Aturia  angustata  (Conrad)  and  a crab  claw. 

Figures  146,  147,  149, 150,  152-154.  Aturia  angustata  (Conrad).  146.  Thin  section  showing  radiating  calcite  within  phragmocone  chamber 
in  crossed-polarized  light,  x70.  LACMP  6699.  147.  Showing  suture,  x0.8;  height  95  mm.  LAM  Loc.  5287,  LACMP  6700.  149.  Showing 
silicified  siphuncular  neck  and  orifice;  height  43  mm.  LACMP  670 1 . 150.  Showing  sutures;  height  65  mm,  width  27  mm  (maximum  diameter). 
Same  specimen  shown  in  Figure  145.  LACMP  6693.  152.  Broken  specimen  showing  siphuncular  orifice  and  neck;  height  1 14  mm.  LACMP 
6702.  153.  Cross  section  showing  funnel-shaped  siphuncular  necks,  x2.0;  length  45  mm.  LACMP  6703.  154.  Broken  specimen  showing 
siphuncular  orifice;  height  75  mm.  LACMP  6704. 

Figures  148,  151.  Crab  claw;  length  40  mm.  LACMP  6705. 


30  Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks 


i \ V- 

, r;?!  a 


132 


137 


143 


144 


145 


Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks  33 


on  the  basis  of  the  size  and  configuration  of  the  burrows,  two 
different  genera  may  be  present. 

SCAPHOPOD 

Dentaliidae 

Denta/ium  ( Fiss ide ntalium 7 ) sp. 
cf.  D.  porter ensis  (Weaver) 

Dentalium  ( Fiss identalium?)  sp.  cf.  D.  porterensis  (Weaver, 
1912:79,  pi.  13,  fig.  1 13)  is  circular  in  cross  section,  slightly 
tapered,  and  has  perhaps  32  (16  exposed  on  one  half)  fine 
radial  riblets  crossed  by  strong  concentric  threads  that  pro- 
duce a basket-weave  sculpture.  The  preservation  does  not 
permit  comparison  with  other  ribbed  Tertiary  dentaliids. 

CEPHALOPODS 

Aturidae 

Aturia  august  at  a (Conrad) 

Figures  133,  135,  137,  140,  143-145,  146, 

147,  149,  150,  152-154,  155-159 

Nautilus  angustatus  Conrad,  1848:728,  pi.  20,  figs.  5,  6. 

Aturia  angustata  has  been  described  in  detail  by  Schenck 
(1931:457-462)  and  Miller  (1947:85-88),  and  the  type  spec- 
imen figured  and  discussed  by  Moore  (1963:85-86,  pi.  31, 
figs.  1,  5). 

A total  of  1 80  specimens  of  A.  angustata  is  in  the  Knappton 
collections,  making  this  cephalopod  by  far  the  most  abundant 
mollusk  collected.  Taking  into  account  a possible  bias  in 
favor  of  collecting  Aturia,  this  is  still  a large  number.  Kummel 
( 1 956:330-33 1 ) called  attention  to  the  rarity  of  post-Triassic 
nautiloids  saying  that  no  large  collection  representing  a pop- 
ulation had  ever  been  assembled  from  a single  horizon  and 
locality.  Stenzel  (in  Ladd,  1957:893)  noted  that  there  are  in 
excess  of  1000  mollusks  representing  other  classes  for  every 
nautiloid  shell  and  that  the  proportion  may  actually  surpass 
10,000  to  1. 

The  shell  of  the  preserved  portion  of  the  living  chamber 
is  commonly  slightly  broken  but  more  frequently  is  intact. 
The  outer  shell  layer  is  dark  brown  and  the  entire  shell  or 
venter  is  thin,  thinner  than  that  of  the  living  Nautilus.  Faint, 
closely  spaced  growth  lines  can  be  seen  on  some  specimens 
(Fig.  135).  The  lateral  lobes  are  tongue-shaped  and  ascending 
on  young  specimens  (Figs.  133,  145,  150),  but  not  on  more 
mature  specimens  (Fig.  147). 

The  specimens  are  believed  to  range  in  size  from  30  to 
180  mm  in  greatest  diameter.  The  smallest  specimens  (25 
mm)  are  not  complete  and  so  were  probably  5 to  10  mm 
larger,  and  the  largest  specimen  measures  170  mm  but  is 


incomplete  and  has  an  estimated  size  of  1 80  mm.  The  largest 
number  of  specimens  sufficiently  complete  to  make  size  mea- 
surement meaningful  (22%)  are  90  mm  in  greatest  diameter. 
Presumably,  this  means  that  many  of  the  specimens  had  not 
reached  maturity  before  death.  About  24%  of  the  specimens 
are  100  to  180  mm  in  maximum  diameter  and  are  assumed 
to  have  been  mature.  The  specimens  are  not  crushed,  and 
none  shows  any  indication  that  it  imploded  as  a result  of 
having  been  transported  to  great  depths. 

The  suture  is  simple  with  a broad  flattened  ventral  saddle, 
a narrow  pointed  lateral  lobe  on  the  umbilical  slope  and 
dorsal  area,  and  a broad  saddle  on  the  dorsal  area  divided 
by  a deep,  narrow  lobe  (Figs.  145,  147,  150).  The  siphuncle 
is  moderate  in  size,  subdorsal  and  marginal  in  position  (Figs. 
137,  140,  143,  144,  149,  152,  154,  155,  157),  and  located 
near  the  apex  in  the  adapical  flexure  of  the  septa.  The  si- 
phuncular  tube  consists  of  a series  of  cone-in-cone  necks,  or 
long  funnel-shaped  connecting  rings  (Fig.  157)  without  the 
long  gaps  between  the  necks  that  are  present  in  Nautilus. 

The  phragmocone  chambers  may  be  filled  with  sediment 
(Fig.  1 54)  but  are  more  commonly  partially  filled  with  calcite 
or  completely  filled  with  calcite,  barite,  quartz,  or  combi- 
nations of  these  minerals  (Fig.  1 53).  A phragmocone  chamber 
of  one  specimen  is  filled  with  glauconite.  Some  of  the  spec- 
imens have  empty  phragmocone  chambers  except  for  a cal- 
cite buttress,  and  these  chambers  may  be  followed  or  pre- 
ceded by  sediment-filled  chambers,  indicating  that  the 
sediment  did  not  enter  through  the  siphuncular  tube  but 
entered  through  a puncture  in  the  shell.  The  body  chamber, 
of  course,  is  always  filled  with  sediment. 

Most  of  the  shells  of  Aturia  angustata  are  preserved  in 
concretions  as  almost  complete  specimens,  but  some  are  frag- 
ments that  may  have  weathered  out  of  concretions  or  not 
have  been  so  preserved.  The  specimens  that  show  a sequence 
of  mineralization,  which  is  currently  being  studied  in  detail, 
begin  with  a buttressing  of  the  shell  walls  with  as  many  as 
nine  layers  of  radial  calcite  (Fig.  146),  followed  by  the  dis- 
solution of  the  aragonitic  shell,  and  then  the  filling  of  the 
shell  cavities  and  the  remaining  chamber  voids  with  calcite, 
barite,  and/or  quartz  in  that  sequence. 

Sepiidae? 

A trace  fossil  that  may  represent  the  cuttlebone  of  a sepiid 
is  illustrated  (Fig.  178). 

FOSSILS  OTHER  THAN  MOLLUSKS 
SPONGES 

Two  sponges  have  been  described  by  J.K.  Rigby  and  D.E. 
Jenkins  (1983)  from  the  upper  part  of  the  Lincoln  Creek 
Formation:  Aphrocallistes  polytretos  (Fig.  180)  and  Eurete 


Figures  155-159.  Aturia  angustata  (Conrad).  155.  Broken  specimen  showing  siphuncular  orifices,  x3;  height  80  mm.  LACMP  6706.  156. 
Showing  phragmocone  chambers  filled  with  wood  fragments  and  sediment.  LACMP  6707. 157.  Silicified  specimen  etched  in  dilute  hydrochloric 
acid  to  show  cone-m-cone,  funnel-shaped  septal  necks,  x 1.5;  47  mm  greatest  diameter.  LACMP  6708.  158.  Fecal  pellets  probably  formed  by 
a manne  worm  in  phragmocone  chamber,  x3.  LACMP  6709.  159.  Fecal  pellets  in  phragmocone  chamber,  x5.  LACMP  6710. 


34  Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks 


Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks  35 


goederti.  In  addition,  hexactinellid  root  tuffs  were  identified 
that  may  represent  a third  sponge.  Although  the  sponges  are 
most  common  stratigraphically  just  below  the  major  mollusk 
locality,  they  also  occur  within  it.  Aphrocallistes  lives  at  depths 
of  100  to  1700  m (Schulze,  1887)  and  Eurete  between  220 
and  715  m,  with  the  majority  of  species  living  at  depths 
between  300  and  360  m (Rigby  and  Jenkins,  1983). 

CORALS 

Dendrophyllia  hannibali  Nomland  (1916:67,  pi.  6,  figs.  1-3) 
was  found  in  one  concretion  that  also  contains  abundant  fish 
debris  and  a small  patch  of  the  siliceous  sponge,  Eurete  goe- 
derti Rigby  and  Jenkins  (1983).  The  specimens  are  poorly 
exposed  and  recrystallized  (Fig.  175),  but  some  show  septa 
(Fig.  176). 

Dendrophyllia  hannibali  was  described  by  Nomland  (1916: 
67)  as  colonial,  branching,  and  forming  several  vertical  series 
that  unite  when  coming  in  contact.  It  has  deep  nearly  round 
calices  with  about  42  to  48  septa.  The  maximum  number  of 
preserved  septa  counted  on  the  specimens  described  by  Nom- 
land is  about  20,  but  recrystallization  is  believed  to  have 
destroyed  many  of  the  septa. 

Dendrophyllia  is  a scleractinian,  ahermatypic  (nonreef- 
building; capable  of  living  in  cold  deep  water)  coral.  Ac- 
cording to  Wells  (1956:F362,  F435),  “the  greatest  develop- 
ment of  ahermatypic  corals  occurs  near  and  down  the  edges 
of  continental  slopes  and  the  equivalent  bathymetric  zone 
around  oceanic  islands  in  depths  from  175  to  about  800  m 
. . . in  temperatures  of  4°  to  21°C.”  Dendrophyllia  is  cos- 
mopolitan in  its  modern  distribution  and  is  known  from  the 
Eocene  through  the  Holocene  at  depths  ranging  from  0 to 
1370  m. 

Flabellum  sp.  (Fig.  129)  has  also  been  collected. 

BRACHIOPOD 

Laqueusl  sp.  cf.  L.  vancouverensis  Davidson  is  poorly  pre- 
served and  only  three  (or  possibly  four)  specimens  have  been 


collected.  The  outer  shell  is  smooth  (Figs.  166,  168,  169),  or 
may  possibly  on  some  specimens  be  finely  ribbed  (Fig.  167), 
and  the  inner  fibrous  layer  is  punctate.  The  specimens  re- 
semble L.  vancouverensis  (Davidson,  1887:1  13,  pi.  18,  figs. 
10-1 3b)  more  than  any  other  described  species.  Gradational 
variation  in  sculpture  between  subspecies  of  the  terebratellids 
(Hertlein  and  Grant,  1 944: 1 32)  seems  to  be  sufficient  to  per- 
haps allow  for  both  smooth-shelled  and  finely  ribbed  forms 
in  one  species. 

ECHINOIDS 

Most  of  the  echinoids  (Figs.  160-162,  164,  170)  are  tests  of 
a spatangoid  (heart  urchin)  that  was  probably  buried  in  living 
position  since  so  many  of  its  spines  are  attached  (Porter  M. 
Kier,  written  commun.,  1980).  The  species  probably  lived 
in  a burrow  at  a depth  of  one  to  several  centimeters  within 
the  sediment.  All  the  tests  are  broken,  perhaps  by  the  weight 
of  the  overburden  as  the  attached  spines  suggest  the  speci- 
mens were  not  transported.  The  echinoids  have  not  been 
found  in  the  center  of  spherical  concretions  typical  of  the  rest 
of  the  fauna. 

A single  specimen  thought  to  be  a madreporite,  a sievelike 
structure  that  provides  access  to  the  water-vascular  system 
(Fig.  163),  was  also  collected. 

LOCALITIES 

NATURAL  HISTORY  MUSEUM  OF  LOS  ANGELES 
COUNTY 

5787.  From  landslide  block  in  upper  part  of  the  Lincoln 
Creek  Formation  between  Knappton  and  Grays  Point,  NW 
'/»  sec.  9,  T.  9 N.,  R.  9 W.,  Knappton  71/2-minute  quadrangle 
(1973  edition),  on  the  Columbia  River,  Washington.  (Gen- 
eral locality  that  includes  5842,  5843,  5844,  and  5852.) 

5802.  From  a limestone  quarry  in  the  siltstones  of  Cliff 
Point  unit  (Wells,  1979)  in  the  bluff  on  the  south  side  of  Bear 
River,  2.3  km  northeast  of  Goulter  Ranch,  on  the  section 
line  between  secs.  20  and  21,  T.  10  N.,  R.  10  W.,  Chinook 


Figures  160-170.  Spatangoid  echinoids  and  a brachiopod. 

Figures  160-162,  164,  170.  Spatangoid  echinoids.  160.  Showing  test  outline  and  spines,  *6.0;  8 mm  greatest  diameter  of  test.  LACMP 
6491.  161.  Showing  ambulacral  area  and  spines,  x3.0.  LACMP  6710.  162.  Showing  test  outline  and  attached  spines,  x3.0;  20  mm  greatest 
diameter  of  test.  LACMP  6492.  164.  Showing  broken  test  with  preserved  ambulacral  areas,  x2.0;  52  mm  greatest  diameter  of  test.  LACMP 
6711.  170.  Showing  outline  of  several  tests  with  associated  wood  fragments.  LACMP  6712. 

Figure  163.  Madreporite?  of  spatangoid  echinoid,  x 12.0;  2.4  mm  greatest  diameter.  LACMP  6713. 

Figures  165-169.  Laqueus ? sp.  cf.  L.  vancouverensis  Davidson.  165.  View  of  apex,  xl.5.  LACMP  6714.  166.  Showing  configuration  and 
narrow  axial  ribs;  length  23  mm,  width  23  mm.  LACMP  6715.  167.  Showing  radial  ribs,  x 1.5;  width  23  mm.  LACMP  6716.  168.  Showing 
configuration;  height  17  mm.  LACMP  6717.  169.  Showing  configuration  and  smooth  shell;  length  35  mm,  width  34  mm.  Same  specimen 
shown  in  Figure  165.  LACMP  6714. 

Figures  171-180.  Teredinid  bores,  crab  claw,  Dendrophyllia  hannibali  Nomland,  a trace  fossil,  and  Aphrocallistes  polytretos  Rigby  and 
Jenkins. 

Figures  171,  172,  177,  179.  Teredinid-bored  wood.  171.  View  of  bored  wood  within  Aturia.  Same  specimen  shown  in  Figure  156,  x3.0. 
LACMP  6707.  177,  179.  Teredinid  tubes  in  wood.  LACMP  6716.  177.  Cross-sectional  view.  179.  Longitudinal  view,  xQ.8. 

Figures  173,  174.  Crab  claw  showing  nodes,  x 1.5;  length  35  mm.  USNM  363992. 

Figures  175,  176.  Dendrophyllia  hannibali  Nomland.  175.  Showing  configuration,  xl.5.  LACMP  6719. 176.  Showing  septa,  x 3.0.  LACMP 
6720. 

Figure  178.  Trace  fossil,  possibly  cuttlebone  of  sepiid,  x 1.5;  length  45  mm.  LACMP  6721. 

Figure  180.  Aphrocallistes  polytretos  Rigby  and  Jenkins.  Showing  irregular  branching  growth.  LACMP  6722. 


36  Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks 


Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks  37 


38  Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks 


7'/2-minute  quadrangle,  about  1 5 km  northwest  of  Knappton, 
Pacific  County,  Washington. 

5842.  From  landslide  block  in  upper  part  of  the  Lincoln 
Creek  Formation  between  Knappton  and  Grays  Point,  in  the 
center  of  the  N 'A,  N V2  sec,  9,  T.  9 N.,  R.  9 W„  Knappton 
7'/2-minute  quadrangle,  on  the  Columbia  River,  Washington. 

5843.  From  landslide  block  in  upper  part  of  the  Lincoln 
Creek  Formation  between  Knappton  and  Grays  Point,  305 
m south  and  430  m east  of  NW  cor.  sec.  9,  T.  9 N.,  R.  9 W., 
Knappton  7 '/2-minute  quadrangle,  on  the  Columbia  River, 
Washington. 

5844.  From  landslide  block  in  upper  part  of  the  Lincoln 
Creek  Formation  between  Knappton  and  Grays  Point,  122 
m east  and  520  m south  of  NW  cor.  sec.  9,  T.  9 N.,  R.  9 W., 
Knappton  7 ‘/2-minute  quadrangle,  on  the  Columbia  River, 
Washington. 

5852.  From  landslide  block  in  upper  part  of  the  Lincoln 
Creek  Formation  between  Knappton  and  Grays  Point,  NE 
'/»,  NW  'A  sec.  9,  T.  9 N.,  R.  9 W.,  Knappton  7 '/2-minute 
quadrangle,  on  the  Columbia  River,  Washington. 

5863.  From  the  Astoria  Formation  between  Knappton  and 
Grays  Point,  SE  'A,  NW  ‘A  sec.  10,  T.  9 N„  R.  9 W„  Knappton 
7'/2-minute  quadrangle,  on  the  Columbia  River,  Washington. 

U.S.  GEOLOGICAL  SURVEY,  MENLO  PARK, 
CALIFORNIA 

M7891.  The  same  locality  as  LAM  5842,  but  collected  by 
E.J.  Moore. 

ACKNOWLEDGMENTS 

I profited  from  discussions  regarding  this  study  with  Kristin 
A.  McDougall  and  George  W.  Moore,  U.S.  Geological  Sur- 
vey, James  C.  Ingle,  Jr.,  Stanford  University,  and  Barry  Roth, 
California  Academy  of  Sciences.  The  constructive  comments 
of  Warren  O.  Addicott  and  George  L.  Kennedy,  U.S.  Geo- 
logical Survey,  are  greatly  appreciated.  The  fossil  photo- 
graphs were  taken  by  Kenji  Sakamoto  and  the  manuscript 
typed  by  Marion  Anderson,  U.S.  Geological  Survey.  I am 
indebted  to  Carole  S.  Hickman,  University  of  California  at 
Berkeley,  and  to  Peter  U.  Rodda,  Barry  Roth,  Robert  Van 
Syoc,  and  Tony  Summers,  California  Academy  of  Sciences, 
for  the  loan  of  specimens.  I am  indebted  to  Edward  C.  Wil- 
son, Los  Angeles  Natural  History  Museum,  for  arranging  the 
loans  of  the  Knappton  collections  and  for  his  support  and 
encouragement  throughout  the  study. 

LITERATURE  CITED 

Abbott,  R.T.  1974.  American  seashells,  2nd  ed.  Van  Nos- 
trand Reinhold,  New  York.  663  pp.,  24  pis. 

Addicott,  W.O.  1970.  Miocene  gastropods  and  biostratig- 
raphy of  the  Kern  River  area,  California.  United  States 
Geological  Survey  Professional  Paper  642:1-174,  21  pis., 
1 1 figs.,  9 tables. 

. 1976a.  Neogene  molluscan  stages  of  Oregon  and 

Washington.  Neogene  Symposium,  Pacific  Section,  So- 
ciety of  Economic  Paleontologists  and  Mineralogists 
Meeting,  San  Francisco,  California.  Pages  95-1 15,5  pis., 
6 figs.,  1 table. 


. 1976b.  New  molluscan  assemblages  from  the  upper 

member  of  the  Twin  River  Formation,  western  Wash- 
ington: significance  in  Neogene  chronostratigraphy. 
United  States  Geological  Survey,  Journal  of  Research 
4(4):437-447,  6 figs.,  2 tables. 

. 1976c.  Molluscan  paleontology  of  the  lower  Mio- 
cene Clallam  Formation,  northwestern  Washington. 
United  States  Geological  Survey  Professional  Paper  976: 
1-44,  9 pis.,  10  figs.,  7 tables. 

Addicott,  W.O.,  Saburo  Kanno,  Kenji  Sakamoto,  and  J.W. 
Miller.  1971.  Clark’s  Tertiary  molluscan  types  from 
the  Yakataga  district  Gulf  of  Alaska.  United  States  Geo- 
logical Survey  Professional  Paper  750-C:  18-33,  6 figs. 

Allison,  R.C.  1978.  Late  Oligocene  through  Pleistocene 
molluscan  faunas  in  the  Gulf  of  Alaska  region.  The  Ve- 
liger  2 1(2):  17 1-1 88,  2 figs.,  2 tables. 

Allison,  R.C.,  and  Louie  Marincovich.  1981.  A late  Oli- 
gocene or  earliest  Miocene  molluscan  fauna  from  Sit- 
kinak  Island,  Alaska.  United  States  Geological  Survey 
Professional  Paper  1233:1-1 1,  3 pis.,  3 figs.,  6 tables. 

Anderson,  F.M.,  and  Bruce  Martin.  1914.  Neocene  record 
in  the  Temblor  Basin,  California,  and  Neocene  deposits 
of  the  San  Juan  district,  San  Luis  Obispo  County.  Cal- 
ifornia Academy  of  Sciences  Proceedings  ser.  4,  3:15- 
1 12,  pis.  1-10. 

Araki,  Yoshio.  1960.  Geology,  paleontology  and  sedimen- 
tary structures  (including  Problematica)  of  the  Tertiary 
formations  developed  in  the  environs  of  Tsu  City,  Mie 
Prefecture,  Japan.  Liberal  Arts  Department,  Mie  Uni- 
versity Bulletin,  Special  Volume  1:1-1 18,  1 1 pis. 

Armentrout,  J.M.  1973.  Molluscan  paleontology  and  stra- 
tigraphy of  the  Lincoln  Creek  Formation,  late  Eocene- 
Oligocene,  southwestern  Washington.  University  of 
Washington,  Seattle,  Ph.D.  dissertation.  479  pp.,  15  pis. 

. 1975.  Molluscan  biostratigraphy  of  the  Lincoln 

Creek  Formation,  southwest  Washington.  Paleogene 
symposium  and  selected  technical  papers.  Pacific  Sec- 
tions American  Association  of  Petroleum  Geologists, 
Society  of  Economic  Paleontologists  and  Mineralogists, 
Society  of  Economic  Geologists,  1975,  Annual  Meeting, 
Long  Beach,  California.  Pages  14-18. 

. 1977.  Cenozoic  molluscan  stages  of  Oregon  and 

Washington.  Geological  Society  of  America,  Abstracts 
with  Programs  9(7):882-883. 

Arnold,  Ralph.  1908.  Description  of  new  Cretaceous  and 
Tertiary  fossils  from  the  Santa  Cruz  Mountains,  Cali- 
fornia. United  States  National  Museum,  Proceedings 
34(  1 6 1 7):345— 390,  pis.  31-37. 

Bernard,  F.R.  1983.  Catalogue  of  the  living  Bivalvia  of  the 
eastern  Pacific  Ocean,  Bering  Strait  to  Cape  Horn.  Ca- 
nadian Special  Publication  of  Fisheries  and  Aquatic  Sci- 
ences 60:102. 

Clark,  B.L.  1918.  The  San  Lorenzo  series  of  middle  Cali- 
fornia. University  of  California  Publications,  Depart- 
ment of  Geology,  Bulletin  1 1(2):45-234,  pis.  3-24. 

. 1925.  Pelecypoda  from  the  marine  Oligocene  of 

western  North  America.  University  of  California  Pub- 
lications, Department  of  Geological  Sciences,  Bulletin 
1 5(4):69— 1 36,  pis.  8-22. 

. 1932.  Fauna  of  the  Poul  and  Yakataga  Formations 


Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks  39 


(upper  Oligocene)  of  southern  Alaska.  Geological  Soci- 
ety of  America,  Bulletin  43(3):797-846,  pis.  14-21. 

Clark,  B.L.,  and  Ralph  Arnold.  1923.  Fauna  of  the  Sooke 
Formation,  Vancouver  Island,  with  description  of  a new 
coral  by  T.  Wayland  Vaughan.  University  of  California 
Publications,  Department  of  Geological  Sciences,  Bul- 
letin 14(5):  123-234,  pis.  15-42. 

Clench,  W.J.,  and  R.D.  Turner.  1950.  The  genera  Sthe- 
norytis,  Cirsotrema,  Acirsa.  Opalia,  and  Amaea  in  the 
western  Atlantic.  Johnsonia  2(29):22 1-248. 

Cochram,  J.K.,  D.M.  Rye,  and  N.H.  Landman.  1981. 
Growth  rate  and  habitat  of  Nautilus  pompilius  inferred 
from  radioactive  and  stable  isotope  studies.  Paleobiol- 
ogy 7:469-480. 

Conrad,  T. A.  1848.  Fossil  shells  from  Tertiary  deposits  on 
Columbia  River,  near  Astoria.  American  Journal  of  Sci- 
ence, ser.  2,  5:432-433,  14  figs. 

. 1849.  Fossils  from  northwestern  America.  In  Dana, 

J.  D.,  United  States  Exploring  Expedition  . . . 1 838— 
1842,  under  the  command  of  Charles  Wilkes  1 0:722— 
728  (appendix),  atlas  pis.  1 7-2 1 . (Text  reprinted  in  Dali, 
1909:152-156.) 

Dali,  W.H.  1887.  Supplementary  notes  on  some  species  of 
mollusks  of  the  Bering  Sea  and  vicinity.  United  States 
National  Museum,  Proceedings  9(57 1 ):297— 309,  pis. 
3,  4. 

. 1891.  Scientific  results  of  explorations  by  the  United 

States  Fish  Commission  Steamer  “Albatross”  (20).  On 
some  new  or  interesting  west  American  shells  obtained 
from  the  dredgings  of  the  United  States  Fish  Commis- 
sion Steamer  “Albatross”  in  1888,  and  from  other 
sources.  United  States  National  Museum,  Proceedings 
14(849):  1 7 3— 191,  pis.  5-7. 

. 1892.  Contributions  to  the  Tertiary  fauna  of  Flor- 
ida, with  especial  reference  to  the  Miocene  silex  beds  of 
Tampa  and  the  Pliocene  beds  of  the  Caloosahatchie  Riv- 
er. Wagner  Free  Institute  of  Science,  Transactions  3(2): 
201-473,  pis.  13-22. 

. 1907.  Descriptions  of  new  species  of  shells,  chiefly 

Buccinidae,  from  the  dredgings  of  the  U.S.S.  “Albatross” 
during  1906,  in  the  northwestern  Pacific,  Bering, 
Okhotsk,  and  Japanese  Seas.  Smithsonian  Miscella- 
neous Collections  50(2)1727:139-173. 

. 1909.  Contributions  to  the  Tertiary  paleontology 

of  the  Pacific  Coast,  I.  The  Miocene  of  Astoria  and  Coos 
Bay,  Oregon.  United  States  Geological  Survey,  Profes- 
sional Paper  59:1-278,  23  pis. 

. 1922.  Fossils  of  the  Olympic  Peninsula  [Washing- 
ton], American  Journal  of  Science  ser.  5,  4:305-314. 

. 1925.  Illustrations  of  unfigured  types  of  shells  in 

the  collection  of  the  United  States  National  Museum: 
United  States  National  Museum,  Proceedings  66:1-41, 
art.  17,  no.  2554,  pis.  1-36. 

Davidson,  Thomas.  1887.  A monograph  of  recent  Brach- 
iopoda.  Transactions  of  the  Linnean  Society  of  London, 
Zoology,  ser.  2,  4(2):  1-248. 

Durham,  J.W.  1937.  Gastropods  of  the  family  Epitoniidae 
from  Mesozoic  and  Cenoizoic  rocks  of  the  west  coast  of 


North  America,  including  one  new  species  by  F.  E.  T urn- 
er  and  one  by  R.  A.  Bramkamp.  Journal  of  Paleontology 
1 1(6):479-5 1 2,  pis.  56,  57. 

. 1 944.  Megafaunal  zones  of  the  Oligocene  of  north- 
western Washington.  University  of  California  Publica- 
tions, Department  of  Geological  Sciences,  Bulletin  27(5): 
101-212,  pis.  13-18,  figs.  1-7. 

Gabb,  W.M.  1866.  Tertiary  invertebrate  fossils.  California 
Geological  Survey,  Palaeontology  2(1)  pt.  2:1-38,  pis. 
1-13. 

. 1869.  Tertiary  invertebrate  fossils.  California  Geo- 
logical Survey,  Paleontology,  Cretaceous  and  Tertiary 
fossils  2(  1 ):65— 1 24,  part  3. 

Grant,  U.S.,  IV,  and  H.R.  Gale.  1931.  Catalogue  of  the 
marine  Pliocene  and  Pleistocene  Mollusca  of  California. 
San  Diego  Society  Natural  History,  Memoir  1:1-1036, 
32  pis.,  15  figs.,  3 tables. 

Ham,  C.K.  1952.  Geology  of  Las  Trampas  Ridge,  Berkeley 
Hills,  California.  California  Division  of  Mines,  Special 
Report  22:1-26,  20  pis. 

Hamada,  Takashi,  Ikuo  Obata,  and  Takashi  Okutani.  1 980. 
Nautilus  macromphalus  in  captivity.  Takai  University 
Press,  Tokyo,  Japan.  80  pp. 

Hayashi,  Tadaichi.  1960.  On  a new  subgenus  and  a new 
species  of  Fulgoraria  from  Japan.  Venus,  Japanese  Jour- 
nal of  Malacology  2 1(1):  1-4,  1 pi. 

Hertlein,  L.G.  1963.  A new  species  of  giant  Lima  from  off 
southern  California  (Mollusca:  Pelecypoda).  California 
Academy  of  Sciences,  Occasional  Paper  40:1-6,  3 figs. 

Hertlein,  L.G.,  and  U.S.  Grant,  IV.  1944.  The  Cenozoic 
Brachiopoda  of  western  North  America.  University  of 
California,  Los  Angeles,  Publications  in  Mathematical 
and  Physical  Sciences  3:1-236,  21  pis.,  34  figs. 

Hickman,  C.S.  1969.  The  Oligocene  marine  molluscan  fau- 
na of  the  Eugene  Formation  in  Oregon.  University  of 
Oregon,  Natural  History  Museum,  Bulletin  16:1-112, 
14  pis. 

. 1980.  Paleogene  marine  gastropods  of  the  Keasey 

Formation  in  Oregon.  Bulletins  of  American  Paleon- 
tology 78(3 10):  1 — 1 12,  10  pis.,  5 figs. 

Howe,  H.V.W.  1922.  The  Miocene  of  Clatsop  and  Lincoln 
Counties,  Oregon.  Stanford  University,  California,  Ph.D. 
dissertation.  286  pp.,  14  pis.,  1 map. 

Javidpour,  Mahdokt.  1973.  Some  records  on  west  Amer- 
ican Cenozoic  gastropods  of  the  genus  Aforia.  The  Ve- 
liger  15(3):  196-205,  1 pl„  2 figs. 

Kanie,  Yasumitsu,  Yoshio  Fukuda,  Hideaki  Nakayama,  Ku- 
nihiro  Seki,  and  Mutsuo  Hattori.  1980.  Implosion  of 
living  Nautilus  under  increased  pressure.  Paleobiology 
6( l):44-47,  3 figs. 

Kanno,  Saburo.  1971.  Tertiary  molluscan  fauna  from  the 
Yakataga  district  and  adjacent  areas  of  southern  Alaska. 
Palaeontological  Society  of  Japan,  Special  Papers  16: 
1-154,  18  pis.,  20  figs. 

Keen,  A.M.  1971.  Sea  shells  of  tropical  west  America,  2nd 
ed.  Stanford  University  Press,  Stanford,  California.  1 064 
pp.,  3305  figs. 

Kira,  Tetsuaki.  1962.  Shells  of  the  western  Pacific  in  color. 


40  Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks 


Hoikusha  Publishing  Company,  Osaka,  Japan.  224  pp., 
72  pis.  (English  edition). 

Korobkov,  I. A.  1949.  O nakhozhdenii  roda  Psephaea  Crosse 
v sredneeotsenovykh  otlozheniakh  sevemogo  Kavkaza 
[On  the  occurrence  of  Psephaea  Crosse  in  middle  Eocene 
sediments  of  the  northern  Caucasus].  Akademia  Nauk 
SSSR,  Doklady  66(4):693-695,  2 figs. 

. 1955.  Mollyuski  srednego  eotsena  sevemogo  Kav- 
kaza i usloviya  ikh  obitaniya.  Leningradskii  Gosudar- 
stvennyi  Universitet,  Uchenye  Zapiski,  Seriya  Geolo- 
gicheskikh  Nauk;  Uchenye  Zapiski,  no.  189,  pp.  1 58— 
230.  Seriya  Geologo-Pochvennykh  Nauk;  Vestnik  6,  Se- 
riya Geologii  i Geografii. 

Kummel,  Bernard.  1956.  Post-Triassic  nautiloid  genera. 
Harvard  University  Museum  of  Comparative  Zoology, 
Bulletin  1 14(7):324-494,  28  pis.,  35  figs. 

Ladd,  H.S.,  editor.  1957.  Treatise  on  marine  ecology  and 
paleoecology.  2,  paleoecology.  Geological  Society  of 
America,  Memoir  67(2):  1-1077. 

Loel,  Wayne,  and  W.H.  Corey.  1932.  The  Vaqueros  For- 
mation, Lower  Miocene  of  California;  [part]  1,  paleon- 
tology. University  of  California  Publications,  Depart- 
ment of  Geological  Sciences,  Bulletin  22(3):3 1-286,  pis. 
4-65. 

Marincovich,  Louie.  1 979  [ 1 980],  Miocene  mollusks  of  the 
Topsy  Formation,  Lituya  District,  Gulf  of  Alaska  Ter- 
tiary province,  Alaska.  United  States  Geological  Survey, 
Professional  Paper  1 1 25-C:  1 — 13,  30  figs. 

Matthews,  J.F.  1955.  Edison  Groves  area  of  Edison  oil  field. 
California  Oil  Fields  4 1 (2):  1 1-14,  1 pi. 

McLean,  J.H.  1971.  A revised  classification  of  the  family 
Turridae,  with  the  proposal  of  new  subfamilies,  genera, 
and  subgenera  from  the  eastern  Pacific.  The  Veliger  1 4(  1 ): 
H4_130,  4 pis.,  1 table. 

Miller,  A.K.  1947.  Tertiary  nautiloids  of  the  Americas. 
Geological  Society  of  America,  Memoir  23:1-234,  100 
pis.,  30  figs.,  1 table. 

Moore,  E.J.  1963  [1964].  Miocene  mollusks  from  the  As- 
toria Formation  in  Oregon.  United  States  Geological 
Survey,  Professional  Paper  419:1-109,  32  pis.,  9 figs.,  3 
tables. 

. 1976.  Oligocene  marine  mollusks  from  the  Pitts- 
burg Bluff  Formation  in  Oregon.  United  States  Geolog- 
ical Survey,  Professional  Paper  922:1-66,  17  pis.,  5 ta- 
bles. 

. 1984.  Middle  Tertiary  molluscan  zones  of  the  Pa- 
cific northwest.  Journal  of  Paleontology  58(3):7 18-737, 
10  figs. 

Noda,  Hiroshi.  1975.  Turciculid  Gastropoda  of  Japan.  Sci- 
ence Reports  of  Tohuku  University,  Sendai,  Ser.  2 (Ge- 
ology) 45(2):51-82,  pis.  9-12,  3 figs.,  2 tables. 

Nomland,  J.O.  1916.  Corals  from  the  Cretaceous  and  Ter- 
tiary of  California  and  Oregon.  University  of  California 
Publications,  Department  of  Geology,  Bulletin  9(5):  59— 
76,  pis.  3-6. 

Powell,  A. W.B.  1942.  The  New  Zealand  Recent  and  fossil 
Mollusca  of  the  family  Turridae,  with  general  notes  on 


turrid  nomenclature  and  systematics.  Auckland  Institute 
and  Museum  Records  2:1-188,  14  pis.,  6 figs. 

. 1969.  The  family  Turridae  in  the  Indo-Pacific,  part 

2,  the  subfamily  Turriculinae.  Indo-Pacific  Mollusca 
2(  1 0):2 1 5—4 15,  pis.  188-324. 

Reagan,  A. B.  1909.  Some  notes  on  the  Olympic  Peninsula, 
Washington.  Kansas  Academy  of  Sciences,  Transactions 
22:131-238,  6 pis. 

Reeve,  Lovell.  1850.  Conchologia  iconica:  or  illustrations 
of  the  shells  of  molluscous  animals,  6 [1849-1851],  Lon- 
don. 

Rehder,  H.A.  1955.  The  genus  Turcicula  Dali.  Malacologi- 
cal  Society,  Proceedings  31:222-225,  1 pi. 

Rigby,  J.K.,  and  D.E.  Jenkins.  1983.  The  Tertiary  sponges 
Aphrocallistes  and  Eurete  from  western  Washington  and 
Oregon.  Natural  History  Museum  of  Los  Angeles  Coun- 
ty, Contributions  in  Science  344:1-13,  23  figs. 

Schenck,  H.G.  1928.  Stratigraphic  relations  of  western  Or- 
egon Oligocene  Formations.  University  of  California 
Publications,  Department  of  Geological  Sciences,  Bul- 
letin 18(1):  1-50,  18  text  figs. 

. 1931.  Cephalopods  of  the  genus  Aturia  from  west- 
ern North  America.  University  of  California  Publica- 
tions, Department  of  Geological  Sciences,  Bulletin  19(19): 
435-490,  pis.  66-78. 

Schulze,  F.E.  1887.  Report  on  the  Hexactinellida  collected 
by  H.M.S.  Challenger  during  the  years  1873-76.  Report 
of  Scientific  Results  of  the  Voyage  of  the  H.M.S.  Chal- 
lenger, Zoology  21:1-513. 

Shikama,  Tokio.  1967.  System  and  evolution  of  Japanese 
fulgorarid  Gastropoda.  Science  Reports  of  the  Yoko- 
hama National  University,  section  II,  13:23-132,  17 
pis.,  26  figs.,  41  tables. 

Steams,  R.E.C.  1891.  Scientific  results  of  explorations  by 
the  United  States  Fish  Commission  steamer  Albatross, 
no.  17.  Descriptions  of  new  west  American  land,  fresh- 
water, and  marine  shells,  with  notes  and  comments. 
United  States  National  Museum,  Proceedings  13:205- 
225,  pis.  15-17. 

Tegland,  N.M.  1931.  Gastropod  genus  Galeodea  in  the 
Oligocene  of  Washington.  University  of  California  Pub- 
lications, Department  of  Geological  Sciences,  Bulletin 
1 9(  1 8): 387— 444.  pis.  59-65. 

. 1933.  The  fauna  of  the  type  Blakeley  upper  Oli- 

gocene  of  Washington.  University  of  California  Publi- 
cations, Department  of  Geological  Sciences,  Bulletin 
23(3): 8 1 — 174,  pis.  2-15. 

Van  Winkle,  K.E.  1918.  Paleontology  of  the  Oligocene  of 
the  Chehalis  Valley,  Washington.  Washington  Univer- 
sity (Seattle)  Publications  in  Geology  l(2):69-97,  pis. 
6-7. 

Vokes,  H.E.  1955.  Notes  on  the  Tertiary  and  Recent  So- 
lemyacidae.  Journal  of  Paleontology  29(3):534-545. 

. 1957.  Miocene  fossils  of  Maryland.  Maryland  De- 
partment of  Geology,  Mines,  and  Water  Resources,  Bul- 
letin 20: 1-85. 

Weaver,  C.E.  1912.  A preliminary  report  on  the  Tertiary 


Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks  41 


paleontology  of  western  Washington.  Washington  Geo- 
logical Survey,  Bulletin  15:1-80,  pis.  1-15. 

. 1942.  Paleontology  of  the  marine  Tertiary  for- 
mations of  Oregon  and  Washington.  University  of 
Washington  [Seattle],  Publication  in  Geology  5(1—3):  1— 
789,  104  pis.  [1943] 

Wells,  J.W.  1956.  Scleractinia.  In  Moore,  R.  C.,  ed.,  Trea- 
tise on  invertebrate  paleontology,  Coelenterata,  part  F. 
Geological  Society  of  America  and  University  of  Kansas 
Press,  Lawrence.  Pages  F328-F444,  figs.  222-243. 

Wells,  R.E.  1979.  Geologic  map  of  the  Cape  Disappoint- 
ment-Naselle  River  area.  Pacific  County,  Washington. 
United  States  Geological  Survey,  Open-File  Report  79- 
389,  1:48,000. 

Wolfe,  E.W.,  and  E.H.  McKee.  1972.  Sedimentary  and 
igneous  rocks  of  the  Grays  River  Quadrangle,  Washing- 


ton. United  States  Geological  Survey,  Bulletin  1335:1— 
70. 

Woodring,  W.P.  1959.  Geology  and  paleontology  of  Canal 
Zone  and  adjoining  parts  of  Panama;  description  of  Ter- 
tiary mollusks  (Gastropods:  Vermetidae  to  Thaididae). 
United  States  Geological  Survey,  Professional  Paper  306- 
B:  147-239,  pis.  24-37. 

Zullo,  V.A.  1982.  Arcoscapellum  Hoek  and  Solidobalanus 
Hoek  (Cirripedia,  Thoracica)  from  the  Paleogene  of  Pa- 
cific County,  Washington,  with  a description  of  a new 
species  of  Arcoscapellum.  Natural  History  Museum  of 
Los  Angeles  County,  Contributions  in  Science  336:1-9, 
18  figs. 

Submitted  28  September  1983;  accepted  17  February  1984. 


42  Contributions  in  Science,  Number  351 


Moore:  Lincoln  Creek  Formation  Mollusks 


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FISHES  OF  THE  GENUS  NANSENIA  (MICROSTOMATIDAE) 
WITH  DESCRIPTIONS  OF  SEVEN  NEW  SPECIES 

Kouichi  Kawaguchi1  and  John  L.  Butler2 


ABSTRACT.  The  microstomatid  fishes  of  the  genus  Nansenia  are 
reviewed  on  a worldwide  basis  and  1 3 valid  species  are  recognized. 
Of  the  10  nominal  taxa  six  species  are  recognized:  N.  Candida  (with 
N.  sanrikuensis  as  a possible  synonym),  N.  groenlandica,  N.  atlan- 
tica,  N.  oblita,  N.  crassa,  and  N.  ardesiaca  (with  N.  tanakai,  N. 
schmitti,  N.  macrolepis,  and  N.  robusta  as  synonyms).  Seven  new 
species  are  described  based  on  specimens  collected  from  the  Atlantic, 
Pacific,  and  Antarctic  oceans.  Original  descriptions  are  presented  for 
N.  ahlstromi,  N.  antarctica,  N.  pelagica,  N.  megalopa,  N.  longicau- 
da,  N.  tenera,  and  N.  tenuicauda.  The  species  of  Nansenia  are  sep- 
arated into  two  groups  based  on  the  number  of  branchiostegal  rays, 
either  three  or  four;  species  of  each  group  are  distinguished  on  the 
basis  of  the  differences  in  vertebral  and  gill  raker  counts,  predorsal 
length  in  percent  of  standard  length,  and  proportional  size  of  the 
caudal  peduncle.  A key  is  presented. 

Five  species,  N.  atlantica,  N.  oblita,  N.  megalopa,  N.  tenera,  and 
N.  tenuicauda  occur  only  in  the  Atlantic;  three,  N.  ahlstromi,  N. 
Candida,  and  N.  crassa,  are  limited  to  the  Pacific.  Nansenia  ardesiaca 
is  widespread  in  the  slope  waters  of  southeast  Asian  seas,  extending 
northward  to  off  southern  Japan  and  westward  to  off  South  Africa. 
Nansenia  groenlandica  occurs  in  the  subarctic  Atlantic  and  subant- 
arctic  eastern  Pacific.  Antarctic  waters  are  inhabited  by  a circumpolar 
species,  N.  antarctica.  Nansenia  pelagica  and  N.  longicauda  are  found 
in  both  the  Atlantic  and  Pacific  oceans;  the  latter  species  restricted 
to  the  Northern  Hemisphere.  Distributional  patterns  of  Nansenia 
are  discussed  in  relation  to  those  of  other  midwater  fishes. 

Six  species  exhibit  allometric  growth  of  head  length,  predorsal 
length,  and  eye  diameter.  Allometry  may  be  an  adaptation  to  me- 
sopelagic  and  benthopelagic  habitats. 

INTRODUCTION 

Fishes  of  the  microstomatid  genus  Nansenia  are  distributed 
widely  in  the  world  ocean.  They  are  found  in  oceanic  and 
coastal  waters  from  the  subarctic  to  the  subantarctic.  Col- 
lections from  both  midwater  and  bottom  trawls  suggest  that 
these  fishes  inhabit  the  epi-  and  mesopelagic  zones  and  also 
the  benthopelagic  zone  of  continental  or  insular  slopes.  In 
areas  of  high  productivity  examples  of  Nansenia  often  exceed 


1.  Ocean  Research  Institute,  University  of  Tokyo. 

2.  Southwest  Fisheries  Center,  National  Marine  Fisheries  Service, 
National  Oceanic  and  Atmospheric  Administration,  La  Jolla. 


200-300  mm  in  standard  length  and  are  sometimes  captured 
in  large  numbers. 

Notwithstanding  the  considerable  numbers  of  adults  and 
juveniles  taken  with  trawls  and  larvae  collected  in  plankton 
nets,  the  species  are  poorly  known.  Cohen  (1958)  examined 
specimens  of  six  of  seven  nominal  species  that  had  been 
referred  to  Nansenia  and  concluded  that  the  paucity  of  ma- 
terial precluded  a critical  revision.  Since  1958  no  compre- 
hensive taxonomic  work  has  been  done  on  the  genus,  al- 
though four  additional  species  have  been  described  (Blache 
and  Rossignol,  1962;  Lavenberg,  1965;  Abe,  1976;  Kana- 
yama  and  Amaoka,  1983).  In  general,  specimens  of  Nansenia 
are  soft-bodied,  poorly  ossified  and  hence  often  collected  in 
poor  condition.  As  a result  the  nominal  species  are  inade- 
quately known  and  a considerable  number  of  species  remain 
undescribed.  Some  species  are  distributed  worldwide,  but  in 
previous  works  geographical  variation  and  dines  were  not 
considered,  and  specimens  from  new  localities  were  named 
as  new  species.  The  degree  of  ossification  is  different,  both 
among  species  and  growth  stages,  which,  combined  with  the 
lack  of  a complete  size  series  for  all  species,  makes  compar- 
ative osteology  difficult.  For  all  these  reasons  it  is  not  possible 
at  this  time  to  present  a comprehensive  definition  of  the 
genus. 

The  present  study  is  a preliminary  one  that  aims  to  di- 
agnose each  species,  both  previously  described  and  ones  de- 
scribed in  this  paper  with  the  objective  of  contributing  in- 
formation that  will  lead  to  a clearer  definition  of  Nansenia 
and  to  the  osteology,  phytogeny,  zoogeography,  early  life  his- 
tory, and  ecology  of  each  species. 

METHODS  AND  MATERIALS 

Measurements  were  taken  with  dividers  and  recorded  to  the 
nearest  0.1  mm.  Unless  otherwise  indicated,  definitions  fol- 
low Hubbs  and  Lagler  ( 1 964)  and  include  the  following:  stan- 
dard length  (SL);  length  of  head  (HL);  depth  of  body  (BD)— 
vertical  through  origin  of  base  of  dorsal  fin;  least  depth  of 
caudal  peduncle  (CPD);  caudal  peduncle  length  (CPL);  di- 
ameter of  eye  (ED)— horizontal  distance  between  opposite 


Contributions  in  Science,  Number  352,  pp.  1-22 
Natural  History  Museum  of  Los  Angeles  County,  1984 


margins  of  socket;  snout  length  (Sn);  interorbital  length  (In- 
tor);  width  of  body  (BW)— width  just  behind  the  head;  pre- 
dorsal length  (Pre  D);  preanal  length  (Pre  A);  preventral  length 
(PreV)— distance  from  tip  of  snout  to  structural  base  of  out- 
ermost ventral  fin  ray.  Size  of  specimen  is  recorded  in  stan- 
dard length,  with  standard  deviation  and  range  in  parenthe- 
ses. Center  of  body  is  the  midpoint  of  standard  length.  If 
allometric  growth  is  indicated  for  a species,  data  are  pre- 
sented for  different  size  groups. 

Numbers  of  vertebrae  and  median  fin  rays  were  counted 
from  radiographs.  The  ultimate  double  rays  of  the  anal  and 
dorsal  fins  were  counted  as  one.  The  short  spine  sometimes 
found  at  the  base  of  the  outermost  pelvic  fin  ray  was  not 
counted.  In  other  counts,  all  elements  were  included.  The 
upturned  ural  centra  were  counted  as  one.  Teeth  were  count- 
ed on  both  sides  of  the  upper  and  lower  jaws.  Frequency 
distributions  of  gill  raker  and  vertebrae  counts  are  shown 
with  the  number  of  specimens  in  parentheses  following  each 
count. 

In  the  materials  section  of  each  species,  catalogue  numbers 
are  followed  by  the  number  of  specimens  and  their  size  range 
in  parentheses.  The  station  number  is  preceded  by  the  cruise 
number  unless  other  wise  indicated.  The  sampling  depth  is 
indicated  by  m and  the  length  of  wire  out  by  mwo.  The 
following  institutional  abbreviations  are  used. 

UBC  Institute  of  Fisheries,  University  of  British  Colom- 
bia, Vancouver,  Canada. 

CAS  California  Academy  of  Sciences,  San  Francisco,  USA. 

FMNH  Field  Museum  of  Natural  History,  Chicago,  USA. 
ISH  Institute  fur  Seefischerei,  Hamburg,  West  Germany. 
KU  Kochi  University,  Japan. 

KYO  Department  of  Fisheries,  Kyoto  University,  Japan. 

LACM  Natural  History  Museum  of  Los  Angeles  County, 
USA. 

OSU  School  of  Oceanography,  Oregon  State  University, 
Corvallis,  USA. 

SIO  Scripps  Institution  of  Oceanography,  University  of 
California,  La  Jolla,  USA. 

SWFC  Southwest  Fisheries  Center,  National  Marine  Fish- 
eries Service,  La  Jolla,  USA. 


UF  Florida  State  Museum,  University  of  Florida, 
Gainesville,  USA. 

USNM  National  Museum  of  Natural  History,  Smithsonian 
Institution,  Washington,  D.C.,  USA. 

ZMUC  Zoologiske  Museum,  University  of  Copenhagen, 
Denmark. 

Genus  Nansenia  Jordan  and  Evermann,  1896 

Nansenia  Jordan  and  Evermann,  1896:528  (type  species  by 
monotypy,  Microstomus  gron/andicus  Reinhardt,  1839;  cf. 
Follet  and  Cohen,  1958). 

Bathymacrops  Gilchrist,  1922:531  (type  species  by  mono- 
typy, Bathymacrops  macrolepis  Gilchrist,  1922). 
Euproserpa  (subgenus)  Fowler,  1934:256  (type  species  by 
original  designation.  Microstoma  schmitti  Fowler,  1934). 

The  definition  of  the  genus  Nansenia  adopted  here  basically 
follows  Chapman  (1942,  1943,  1948)  and  Cohen  (1964).  The 
placement  of  the  genus  Nansenia  in  the  family  Microstom- 
idae  follows  Ahlstrom  et  al.,  1984.  Body  slender,  subcylin- 
drical  or  slightly  compressed.  Dorsal  adipose  fin  present  on 
the  last  quarter  of  body.  Predorsal  length  less  than  6 1 percent 
of  SL.  Snout  shorter  than  half  of  eye,  which  is  directed  lat- 
erally and  is  not  tubular.  Branchiostegals  three  or  four.  Anal 
fin  rays  8-10  (rarely  11).  Pectoral  fins  inserted  on  sides  of 
the  body.  Parietals  large  and  meeting  on  the  midline.  No 
teeth  on  premaxillary  or  maxillary.  Teeth  present  on  pala- 
tines, head  of  vomer  and  dentaries. 

REMARKS.  The  difference  between  Nansenia  and  Bathy- 
lagus  (presently  placed  in  a separate  family,  Bathylagidae)  is 
not  clear,  despite  efforts  to  discover  adequate  diagnostic  char- 
acters. Characters  concerned  with  the  swimbladder,  the  oto- 
liths, and  morphology  of  the  larvae,  which  may  be  diagnostic, 
are  not  always  known  for  all  members  of  both  genera  at 
different  growth  stages.  Their  value  for  distinguishing  the  two 
genera  remains  to  be  confirmed. 

The  arrangement  of  the  parietals,  which  have  been  re- 
ported to  meet  broadly  on  the  midline  in  Nansenia,  but  not 
meeting  in  Bathylagus  (Chapman,  1942,  1943,  1948;  Cohen, 
1964),  is  probably  diagnostic  if  it  is  partly  modified  to  pa- 


Table  1.  Counts  and  measurements  used  in  the  key  to  the  species  with  three  branchiostegal  rays  and  distributional  information. 


Species 

Vertebrae 

Gill 

rakers 

Predorsal 
distance 
(%  SL) 

Ratio 

CPL/CPD1 

Distribution 

N.  ahlstromi 

35-36 

40 

50.5-55.8 

1.0-1. 3 

Subtropical  Eastern  Pacific,  20°N-35°N 

N.  Candida 

44-47 

26-31 

49.4-57.6 

1. 8-2.2 

Subarctic  Pacific,  30°N-55°N 

N.  groenlandica 

42-45 

37-45 

43.7-45.6 

2. 8-4.0 

Subarctic  Atlantic,  40°N-70°N; 

Eastern  South  Atlantic 

N.  antarctica 

49-50 

35-41 

45.9-47.7 

2.0-2. 5* 

Southern  oceans,  40°S-60°S 

2. 5-3.0** 

1 Caudal  peduncle  length  to  caudal  peduncle  depth. 
* <1 10  mm  SL. 

**>110  mm  SL. 


2 Contributions  in  Science,  Number  352 


Kawaguchi  and  Butler:  Fishes  of  the  Genus  Nansenia 


Table  2.  Counts  and  measurements  used  in  the  key  to  the  species  with  four  branchiostegal  rays  and  distributional  information. 


Species 

Vertebrae 

Gill 

rakers 

Predorsal 

length 

(%  SL) 

Caudal 
peduncle 
length 
(%  SL) 

Caudal 

peduncle 

length 

(%  SL) 

Ratio 

CPL/CPD1 

Distribution 

N.  pelagica 

38-39 

36-41 

50.4-57.8 

1 1.5-13.3 

8.1-1 1.3 

1.1-1. 5 

Tropical  and  Subtropical 
Atlantic  and  Pacific 

N.  atlantica 

41-42 

30-36 

51.9-56.9 

1 1.3-12.2 

8. 2-8. 6 

1.4-1. 5 

Tropical  Atlantic 

N.  oblita 

42-45 

30-36 

55.0-56.1 

9.3-12.2 

8. 2-9. 6 

1. 1-1.2 

Mediterranean  and  Eastern 
North  Atlantic 

N.  crassa 

43-46 

35-37 

49.0-60.2 

13.0-15.2 

6. 3-7. 7 

1. 7-2.2 

Eastern  Tropical  and 
Subtropical  Pacific 

N.  megalopa 

44-45 

21-23 

42.1-44.2 

15.6-17.6 

5.2-5. 7 

2.8-3. 2 

Tropical  Atlantic 

N.  longicauda 

47-50 

23-27 

41.8-46.0 

15.4-18.6 

4.3-5. 6 

3.0-4. 1 

North  Atlantic  and  North 
Pacific,  20°N-40°N 

N.  tenera 

42-43 

43-46 

43.4-45.8 

12.6-15.0 

6. 2-6.5 

1. 9-2.4 

Subpolar  and  Temperate 
Atlantic 

N.  ardesiaca 

46-48 

27-35 

45.7-48.5 

13.0-15.0 

6. 1-7.5 

1. 7-2.3 

Off  Japan,  Southeast 
Asia  and  East  Africa 

N.  tenuicauda 

46 

38-42 

44.0-45.8 

13.5-13.9 

4. 4-5.0 

2.7-3. 1 

South  Atlantic 

1 Caudal  peduncle  length  to  caudal  peduncle  depth. 


rietals  of  Bathylagus  not  meeting  or  meeting  at  a point  in 
the  anteriormost  part  on  midline.  Branchiostegal  counts  are 
three  or  four  in  Nansenia,  but  mostly  two  in  Bathylagus. 
However  at  least  one  undescribed  bathylagid  examined  dur- 
ing the  present  study  had  exceptionally  three  branchiostegals. 
Anal  fin  ray  counts  are  also  useful  in  separating  the  two 
genera:  8-10  (rarely  1 1)  in  Nansenia  and  12-28  (rarely  1 1) 
in  Bathylagus.  Better  diagnostic  characters  for  the  genus  Nan- 
senia must  await  a revision  of  the  family  Bathylagidae. 

KEY  TO  SPECIES 

The  species  of  Nansenia  are  separated  into  two  groups  based 
on  branchiostegal  counts  of  three  or  four.  Four  species  have 
three  branchiostegals  and  nine  have  four. 

Counts  and  measurements  used  in  the  keys  and  distribu- 
tional data  are  presented  for  all  species  in  Tables  1 and  2 
and  Figures  21  and  22.  After  identification  is  made  with  the 
key,  specimens  should  be  checked  against  the  information 
in  the  tables  and  also  against  the  detailed  descriptions  of  each 
species.  The  number  of  branchiostegal  rays  of  Nansenia  is 
stable  within  species,  but  careful  examination  under  a dis- 
secting microscope  is  necessary  to  recognize  the  innermost 
small  ray  (Fig.  1). 

KEY  TO  SPECIES  OF  NANSENIA  WITH  THREE 
BRANCHIOSTEGAL  RAYS 


1 a.  Ratio  of  caudal  peduncle  length  to  caudal  peduncle  depth 

less  than  1.5;  vertebrae  35-36  ahlstromi 

1 b.  Ratio  of  caudal  peduncle  length  to  caudal  peduncle  depth 
greater  than  1.5;  vertebrae  40-42  2 


2a.  Gill  rakers  on  first  arch  25-31;  predorsal  length  more 

than  49  percent  of  SL Candida 

2b.  Gill  rakers  on  first  arch  35-45,  predorsal  length  less  than 
49  percent  of  SL  in  specimens  larger  than  50  mm  SL  . 

3 

3a.  Vertebrae  42  (eastern  South  Pacific),  43-45  (Atlantic) 

groenlandica 

3b.  Vertebrae  49-50  antarctica 

KEY  TO  SPECIES  OF  NANSENIA  WITH  FOUR 
BRANCHIOSTEGAL  RAYS 

la.  Vertebrae  38-39  pelagica 

lb.  Vertebrae  41-50  2 

2a.  Predorsal  length  49  or  more  percent  of  SL  3 

2b.  Predorsal  length  less  than  49  percent  of  SL  5 

3a.  Length  of  caudal  peduncle  more  than  12.5  percent  of  SL 

and  depth  of  caudal  peduncle  less  than  8 percent  of  SL 

with  CPL/CPD  ratio  more  than  1.6  crassa 

3b.  Length  of  caudal  peduncle  less  than  12.5  percent  of  SL 
and  depth  of  caudal  peduncle  more  than  8 percent  of  SL 

with  CPL/CPD  ratio  less  than  1.6  4 

4a.  Proximal  part  of  adipose  fin  densely  pigmented;  verte- 
brae 41-42;  gill  raker  count  30-36  atlantica 

4b.  Proximal  part  of  adipose  fin  not  pigmented;  vertebrae 

42-45;  gill  raker  count  28-30  oblita 

5a.  Length  of  caudal  peduncle  more  than  1 5.2  percent  of  SL 
and  depth  of  caudal  peduncle  less  than  6 percent  of  SL; 

gill  rakers  21-27  6 

5b.  Length  of  caudal  peduncle  less  than  15.2  percent  of  SL 
and  depth  of  caudal  peduncle  more  than  6 percent  of  SL 
except  in  N.  tenuicauda  7 


Contributions  in  Science,  Number  352 


Kawaguchi  and  Butler:  Fishes  of  the  Genus  Nansenia  3 


a 


b 


Figure  1.  Typical  patterns  of  branchiostegal  rays  of  Nansenia.  a. 
N.  ardesiaca,  N.  ahlstromi,  N.  crassa,  and  N.  megalopa ; b.  N.  pe- 
lagica  and  N.  atlantica;  c.  N.  tenera  and  N.  longicauda;  d.  N.  ant- 
arctica;  e.  N.  groenlandica. 


6a.  Vertebrae  44-45;  gill  rakers  2 1-23  megalopa 

6b.  Vertebrae  47-50;  gill  rakers  23-27  longicauda 

7a.  Vertebrae  42-43;  gill  rakers  43-46  tenera 

7b.  Vertebrae  46-48;  gill  rakers  27-42  8 


8a.  Gill  rakers  27-35;  CPL/CPD  ratio  less  than  2.5 

ardesiaca 

8b.  Gill  rakers  38-42;  CPL/CPD  ratio  more  than  2.5  .... 
tenuicauda 

SPECIES  WITH  THREE  BRANCHIOSTEGAL  RAYS 

Nansenia  ahlstromi  new  species 

Figures  2,  21 

HOLOTYPE.  SIO  57-87  (1 , ca.  95  mm,  slightly  damaged 
mature  female),  R/V  Spencer  F.  Baird,  29°15'N,  126°07'W, 
2240-0730,  14-15  May  1955,  10  ft.-IKMT,  0-754  m. 

PARATYPES.  SIO  63-425  (1,  38.7),  R/V  Horizon, 
27°46'N,  1 29°14.9'W,  0534-1006,  2 Apr.  1962,  10  ft.- 
IKMT,  4500  mwo;  LACM  43546-1  (1,  32.0),  R/V  David 
Starr  Jordan,  24°00'N,  145°00'W,  2348-0021,  21-22  May 
1972,  6 ft.-IKMT  600  mwo;  SWFC  7210-24.139  (1,  58.5 
decomposed),  R/V  David  Starr  Jordan,  24°00'N,  139°00'W, 
0040-0159,  29  Oct.  1972,  50  ft.  universal  MWT,  0-494  m. 

DIAGNOSIS.  Differs  from  other  Nansenia  in  having  three 
branchiostegal  rays  and  35-36  vertebrae,  the  least  of  any 
known  species. 

DESCRIPTION.  Counts  and  measurements  are  based  on 
three  juvenile  specimens,  32.0-58.5  mm  SL.  Only  the  ver- 
tebrae were  counted  for  the  damaged  adult  specimen,  ca.  95 
mm  SL.  D 8-9;  A 7-8;  P 12;  V 9-1 1;  gill  rakers  12  + 28  in 
two  specimens  and  11  + 29  in  one,  total  40;  branchiostegal 
rays  three;  vertebrae  35  in  one  specimen,  36  in  three  speci- 
mens. Pyloric  caeca  not  counted  due  to  paucity  of  specimens. 
About  30  conical  teeth  irregularly  arranged  on  head  of  vomer; 
about  100  compressed  teeth  closely  set  on  dentaries  in  ho- 
lotype. 

Head  relatively  large  compared  with  body,  reflecting  the 
reduced  number  of  vertebrae.  Eye  large,  diameter  2. 3-2. 9 
times  in  head  length  in  three  specimens,  38.7-ca.  95  mm  SL. 


Head  length  and  eye  diameter  indicate  remarkable  allometric 
growth  (Fig.  4).  Supraorbital  bone  damaged  and  lost  in  the 
holotype.  Pectoral  fin  base  low,  its  upper  end  above  ventral 
margin  of  body  about  one-third  of  the  distance  between  ven- 
tral margin  and  lateral  line.  Origin  of  dorsal  fin  base  behind 
center  of  body.  Ventral  fin  base  below  or  just  behind  posterior 
end  of  dorsal  fin  base  in  the  three  juveniles,  32.0-58.5  mm 
SL,  but  well  behind  dorsal  fin  base  in  the  holotype.  Adipose 
fin  base  over  posterior  half  of  the  anal  fin  base. 

Tip  of  lower  jaw,  anterior  part  of  gular  area,  and  base  of 
adipose  fin  darkly  pigmented  in  the  38.7-mm  SL  specimen 
(SIO  63-425).  The  same  pigment  pattern  is  present  in  the 
holotype. 

SIZE.  The  holotype  of  ca.  95  mm  is  an  adult  bearing  eggs. 

DISTRIBUTION.  Restricted  to  the  subtropical  eastern 
North  Pacific. 

ETYMOLOGY.  It  is  with  great  pleasure  that  we  name  this 
species  Nansenia  ahlstromi  in  recognition  of  the  contribu- 
tions of  the  late  Elbert  Halvor  Ahlstrom  to  our  knowledge 
of  pelagic  fishes. 

Nansenia  Candida  Cohen,  1958 

Figures  3,  4,  22 

Nansenia  Candida  Cohen,  1958:52-54,  fig.  1 (orig.  descr.) 

Eastern  North  Pacific. 

INansenia  sanrikuensis  Kanayama  and  Amaoka,  1 983:77— 

79  (orig.  descr.)  Western  North  Pacific. 

MATERIALS.  CAS  51023  (1,  alizarin  specimen  dissect- 
ed), one  of  the  paratypes,  R/V  Hugh  M.  Smith,  41°39'N, 
139°02'W,  30  May  1956,  from  the  stomach  of  Alepisaurus 
sp.;  OSU  8,  9 (2,  46.4,  80.7),  49°16'N,  132°42'W,  0-60  m; 
OSU  10,  12  (2,  44.0,  136.9),  41°59.5'N,  126°30.6'W,  0-200 
m;  OSU  11  (1,  150.5),  44°37.3'N,  125°17.3'W,  1500-500  m; 
OSU  960  (1,  150.5),  44°37.3'N,  125°38.0'W,  0-200  m;  OSU 
1132,1133  (2,  79.0,  72.2),  44°25'N,  1 29°35'W,  0-75  m;  OSU 
1 166  (1,  32.1),  MT-648,  44°16'N,  125°15'W,  surface  tow,  40 
mwo;  OSU  1 170  (1,23.5),  MT-652,  44°3  l'N,  125°17'W,  0- 
1400  m;  OSU  1191  (2,  95.8,  99.6),  MT-757,  44°38'N, 
1 28°35'W,  0-210  m;  OSU  1212  (1,  140.0),  44°39.8'N, 
125°00'W,  0-200  m;  OSU  1216  (1,  108.2),  MT-749,  44°39'N, 
1 25°36'W,  0-200  m;  OSU  1 2 1 7 ( 1 , 60.3),  MT-727,  44°39'N, 
1 28°00'W,  0-1000  m;  OSU  2003  (1,  179.2),  haul  2054, 
44°49.7'N,  1 25°34.0'W,  0-600  m;  OSU  2388  ( 1 , 1 29. 1 ),  MT- 
2398,  44°39'N,  125°29'W,  0-500  m;  USNM  197380  (1,  60.5), 
20  miles  southeast  of  San  Clemente  Is.,  California,  spit  up 
by  albacore;  USNM  195874  (1,  122),  sta.  BB-76,  North  Pa- 
cific, 0-30  m;  USNM  195875  (1,  75.5),  sta.  BB- 1 76,  48°03'N, 
1 34°20'W;  USNM  195876  (1,  156),  sta.  BB-199,  49°15.6'N, 
145°56.7'W,  100  mwo;  SIO  66-5 1-9F  (1,  24),  40°35'N, 
125°51.5'W;  SIO  55-73-9A  (1,  ca.  64,  damaged),  39°01'N, 
165°10'W,  from  Alepisaurus  stomach;  UBC  65-607  (2,  75.5, 

81.0)  and  UBC  65-609  (2,  76.5,  87.5),  off  Queen  Charlotte 
Sound,  B.C.,  Canada;  UBC  65-6 10  ( 1 6,  76.0-88.0)  and  UBC 
65-615  (1,  77.5),  off  Queen  Charlotte  Sound;  UBC  65-623 
(16,  118.0-187.0),  52°11'N,  133°11'W;  UBC  65-524  (1, 

121.0) ,  52°1 3'N,  1 33°1 2'W;  LACM  34258-1  (2,  63-71), 


4 Contributions  in  Science,  Number  352 


Kawaguchi  and  Butler:  Fishes  of  the  Genus  Nansenia 


h 


■*  1 0 mm 


Figure  2.  Nansenia  ahlstromi.  Paratype,  SIO  63-425,  38.7  mm  SL. 

R/V  Alaska,  off  San  Clemente  Is.,  California;  LACM  9006- 
23  (1,  35),  R/V  Velero,  Catalina  Basin  off  California. 

DIAGNOSIS.  Differs  from  other  Nansenia  in  the  follow- 
ing combination  of  characters,  three  branchiostegal  rays  and 
26-31  gill  rakers  on  the  first  arch.  Vertebral  counts  higher 
(44-47)  in  N.  Candida  than  in  N.  ahlstromi  (35-36). 

DESCRIPTION.  D 9-10;  A 8-9;  P 10-1 1 (rarely  9);  V 9- 
10  (rarely  1 1);  gill  rakers  8-1 1 + 1 + 17-20,  total  26-3 1 with 
26  in  one  specimen,  27  in  five,  28  and  29  in  three  respectively, 


30  in  one  and  31  in  three;  branchiostegal  rays  3;  vertebrae 
44-47  with  44  in  two  specimens,  45  in  five,  46  in  eight  and 
47  in  two.  The  above  counts  are  based  on  18  specimens, 
46.2-179.2  mm  SL.  Pyloric  caeca  seven  in  two  specimens, 
nine  in  one.  About  12-21  conical  teeth  on  head  of  vomer, 
about  80-100  teeth  on  dentaries,  resembling  a closely  spaced 
picket  fence,  in  three  specimens  more  than  130  mm  SL. 

Body  slender,  moderately  compressed,  with  body  depth  to 
width  ratio  1.5-1. 7.  Eye  diameter  3.0  to  3.4  times  in  head 


Figure  3.  Nansenia  Candida.  OSU  2003,  mature  female,  179.2  mm  SL. 


Contributions  in  Science,  Number  352 


Kawaguchi  and  Butler:  Fishes  of  the  Genus  Nansenia  5 


% 

70 


PreD 


(D 

O 

0) 

0. 


40- 


30- 


HL 


20-  * 


o 


0 ' ’ ' >- 

0 50  100  150  200 


S L (mm) 


Figure  4.  Nansenia  Candida.  Predorsal,  head  lengths,  and  eye  di- 
ameter in  relation  to  standard  length.  Triangles  indicate  the  values 
for  postlarvae  and  young  juveniles  smaller  than  23  mm  SL. 


length,  viewed  laterally,  its  dorsal  margin  (supraorbital  bone) 
touching  or  slightly  higher  than  dorsal  margin  of  head,  yel- 
lowish iridescent  pigment,  roughly  crescent-shaped,  present 
in  the  posterior  half  of  iris.  Aphakic  space  developed,  its 
horizontal  space  nearly  equal  to  width  of  anteriormost  part 
of  iris,  and  about  half  the  width  of  its  posterior  part.  Aphakic 
space  less  well  developed  in  juveniles  smaller  than  100  mm 
SL.  Allometric  growth  in  head  length,  eye  diameter  and  pre- 
dorsal length  is  shown  in  Figure  4. 

Upper  end  of  pectoral  fin  base  on  midpoint  between  lateral 


line  and  ventral  margin  of  body,  or  slightly  lower;  its  lower 
end  separated  from  ventral  margin  of  body  by  a distance 
more  than  length  of  pectoral  fin  base.  Dorsal  fin  originating 
behind  center  of  body.  Ventral  fin  base  below  posterior  part 
of  dorsal  fin.  Origin  of  anal  fin  slightly  in  advance  of  adipose 
fin.  Skin  around  the  anterior  end  of  adipose  and  caudal  fin 
bases  densely  pigmented.  Body  color  in  formalin  preserved 
specimens  brown.  However,  Cohen  (1958)  noted  that  in  life 
the  entire  fish  is  probably  a bright  silver. 

SIZE.  The  largest  specimen  examined,  179.2  mm  SL,  had 
mature  ovaries. 

DISTRIBUTION.  Nansenia  Candida  has  been  collected 
in  the  subarctic  eastern  Pacific  east  of  170°W  between  35°N 
and  55°N  and  in  the  California  Current  region  north  of  30°N 
(Fig.  22). 

REMARKS.  Kanayama  and  Amaoka  (1983)  described 
Nansenia  sanrikuensis  based  on  two  large  adult  specimens, 
206.0  and  238.8  mm  SL.  This  species  is  closely  related  to 
N.  Candida,  and  both  species  occur  in  the  subarctic  North 
Pacific.  But  N.  Candida  has  been  reported  only  from  the 
eastern  part  of  the  Pacific  between  the  latitude  of  southern 
California  and  Canada,  and  N.  sanrikuensis  was  collected  in 
the  western  Pacific  off  northern  Honshu,  Japan.  Kanayama 
and  Amaoka  (1983)  reported  that  N.  sanrikuensis  differs  in 
having  a smaller  eye  (4. 8-5. 8 vs.  8. 3-9. 6 percent  of  SL), 
shorter  snout  (3. 5-3.6  vs.  5. 1-5.8),  lower  gill  raker  counts 
(25-27  vs.  30)  and  higher  vertebral  counts  (48-49  vs.  44- 
47).  Their  proportional  data  and  gill  raker  counts  for  N. 
Candida  are  based  on  the  original  description  by  Cohen  ( 1 958) 
which  was  taken  from  three  specimens,  54.4-72.4  mm  SL, 


Figure  5.  Nansenia  groenlandica.  ISH  194-59,  1 12.5  mm  SL. 


6 Contributions  in  Science,  Number  352 


Kawaguchi  and  Butler:  Fishes  of  the  Genus  Nansenia 


for  measurements  and  five  for  meristic  counts.  However, 
their  vertebral  counts  are  based  on  the  present  study.  Spec- 
imens studied  by  Kanayama  and  Amaoka  ( 1 983)  and  Cohen 
(1958)  differ  greatly  in  size.  In  the  present  study  we  found 
considerable  allometric  growth  in  eye  diameter,  snout  length, 
and  head  length  of  N.  Candida.  We  conclude  that  proportions 
of  snout,  eye,  and  head  are  invalid  for  distinguishing  the  two 
species.  The  ranges  of  gill  raker  counts  are  also  overlapping 
between  the  two  (25-27  vs.  26-31)  as  N.  Candida  has  a 
wider  range  than  Cohen  (1958)  described.  At  present  only 
vertebral  counts  (48-49  vs.  44-47)  separate  these  species. 
Considering  the  adjacent  ranges,  the  paucity  of  specimens  of 
N.  sanrikuensis  and  the  possibility  of  a geographical  cline, 
the  status  of  N.  sanrikuensis  is  questionable. 


% 

50 


40 


30 


20  - 


10 


Pre  D 

~o  °~o 


HL 


~o  O 7T°~ 


ED 


Nansenia  groenlandica  (Reinhardt,  1840) 

Figures  5,  6,  21 


0 I— 

40 


60 


80 


100 


S L |mm| 


— i -i — 

120  140 


Microstomas  gronlandicus  Reinhardt,  1840:8  (orig.  descr. 

Fiskenesset,  Greenland). 

Nansenia  groenlandica:  Jordan  and  Evermann,  1896:528; 

Schmidt,  1918:12,  figs.  1-12;  Cohen,  1964:24,  fig.  8. 

MATERIALS.  Atlantic:  ISH  21/75  (2,  57.0-67.0),  FFS 
Walther  Herwig  sta.  10-1/75,  41°18'N,  27°49'W,  0-117  m; 
ISH  29/75  (1,  32.5),  FFS  Walther  Herwig  sta.  14-1/75, 
41°26'N,  27°09'W,  0-183  m;  ISH  194/59  (1,  112.5),  FFS 
Anton  Dorn  sta.  3257/59,  6 1°48'N,  1 3°08'W,  0-1400  m;  ISH 
314/73  (2,  132.0,  damaged),  FFS  Walther  Herwig  sta.  678/ 
73,  65°09'N,  32°50'W,  0-1 100  m;  ISH  448/73  (8,  60.5-103.5), 
FFS  Walther  Herwig  sta.  693/73,  57°55'N,  28°23'W,  0-415 
m;  ISH  473/73  (1,  89.0),  FFS  Walther  Herwig  sta.  693/73, 
56°28'N,  26°44'W,  0-2500  m;  ISH  540/73  (3,  80.0-108.0), 
FFS  Walther  Herwig  sta.  696/73,  55°39'N,  25°47'W,  0-2500 
m;  ISH  540/73  (3,  80.0-108.0),  FFS  Walther  Herwig  696/ 
73,  55°39'N,  25°47'W,  0-410  m;  ISH  748/73  (1,  57.0),  FFS 
Walther  Herwig  707/73,  50°03'N,  19°39'W,  0-255  m;  USNM 
186071  (damaged,  head  only),  M/V  Delaware.  40°34'N, 
64°07'W,  from  stomach  of  Makaira\  USNM  221368  (3,  dam- 
aged, ca.  63-84),  FFS  Walther  Herwig  sta.  695/73,  55°43'N, 
25°53'W,  0-2600  m.  Pacific:  LACM  1 1292  (1,  63.7),  R/V 
Eltanin  sta.  1972,  39°34'S,  127°18'W,  IKMT,  0-4493  m; 
LACM  10288  (2,  damaged,  ca.  30),  R/V  Eltanin  sta.  190, 
34°5  l'S,  74°04'W,  0-1580  m. 

DIAGNOSIS.  Differs  from  other  Nansenia  in  having  three 
branchiostegal  rays,  42-45  vertebrae  (43-45  in  the  Atlantic, 
42  in  Pacific  specimens),  and  37-45  gill  rakers  on  the  first 
arch. 

DESCRIPTION.  D 9-10;  A 8-10;  P 1 1-13;  V 1 l-12(rare- 
ly  10);  gill  rakers  12-15  + 1 + 23-29,  total  37-45  with  37(2), 
38(5),  40(1),  41(2),  42(1),  44(2),  and  45(1);  branchiostegal 
rays  three;  vertebrae  42(1)  in  the  South  Pacific,  43-45  with 
43(8),  44(9),  and  45(1)  in  the  North  Atlantic.  Counts  are 
based  on  19  specimens,  48.0  to  132.0  mm  SL.  Pyloric  caeca 
seven  and  eight  in  two  specimens;  20-22  conical  teeth  irreg- 
ularly arranged  on  vomer,  50-65  teeth  closely  set  on  dentaries 
in  four  specimens. 

Body  slender,  body  depth-width  ratio  ranges  from  1.0- 
1.2.  Eye  large,  its  diameter  2.5  to  2.8  times  in  head  length. 


Figure  6.  Nansenia  groenlandica.  Predorsal,  head  lengths,  and  eye 
diameter  in  relation  to  standard  length. 


Viewed  laterally,  about  one-third  of  eye  circumference  pro- 
truded above  the  dorsal  margin  of  the  head.  White  or  yel- 
lowish tissue  developed  over  posterior  half  of  iris,  sometimes 
covering  whole  iris.  Aphakic  space  slightly  developed  ante- 
riorly, its  horizontal  space  nearly  equal  to  half  to  two-th(rds 
width  of  the  anterior  part  of  iris.  Aphakic  space  less  well 
developed  in  juveniles  smaller  than  100  mm  SL. 

Upper  end  of  pectoral  fin  base  separated  from  ventral  mar- 
gin of  body  by  one-third  of  distance  between  lateral  line  and 
ventral  margin  or  slightly  higher;  its  lower  end  above  ventral 
margin  by  a space  more  than  the  length  of  pectoral  fin  base. 
Dorsal  fin  originating  in  front  of  center  of  body.  Ventral  fin 
base  below  posterior  end  of  dorsal  fin  base  or  just  behind  it. 
Adipose  fin  base  above  last  two  anal  fin  rays.  No  remarkable 
inflection  in  allometric  growth  for  specimens  between  48  and 
132  mm  SL  (Fig.  6). 

Skin  around  adipose  fin  base  uniformly  pigmented  com- 
pared to  the  lateral  body  surface.  Caudal  fin  base  and  pos- 
terior part  of  caudal  peduncle  densely  pigmented,  sparser  in 
specimens  larger  than  100  mm  SL.  Body  of  preserved  spec- 
imens usually  brown  or  light  brown,  some  with  remains  of 
guanine.  The  entire  body  surface  of  the  ISH  1 94-59  specimen, 
although  all  scales  had  come  off,  is  covered  with  guanine. 
This  specimen  may  have  been  preserved  in  alchohol. 

SIZE.  The  badly  damaged  specimen  (caudal  segment  miss- 
ing) taken  from  a marlin  stomach,  USNM  1 8607 1 , identified 
as  N.  groenlandica  only  by  its  branchiostegal  ray  count  and 
locality,  is  a mature  male  and  its  standard  length  can  be 
estimated  as  about  180  mm  based  on  its  head  length.  The 
largest  four  intact  specimens,  103.0  to  132.0  mm,  collected 
in  September  have  undeveloped  gonads. 

DISTRIBLTTION.  This  species  occurs  widely  in  the  arctic 
and  subarctic  waters  of  the  North  Atlantic  Ocean.  The  south- 
ern limit  of  its  distribution  is  near  40°N.  Present  results  agree 
well  with  the  work  of  Schmidt  (1918)  in  the  eastern  North 
Atlantic.  Three  specimens  collected  in  the  subantarctic  or 


Contributions  in  Science,  Number  352 


Kawaguchi  and  Butler:  Fishes  of  the  Genus  Nansenia  7 


CM  I 
0 


23456789  10 


I I I I CM 
12  13  14  15 


Figure  7.  Nansenia  antarctica.  LACM  10875  (one  of  the  three  catalogued),  198.9  mm  SL,  holotype. 


transitional  waters  of  the  eastern  South  Pacific  show  no  mor- 
phological difference  from  the  Atlantic  specimens,  but  have 
slightly  fewer  vertebrae.  Mukhacheva  (1972)  reported  a sim- 
ilar distribution  pattern  for  the  midwater  fish,  Gonostoma 
bathyphilum,  which  is  widely  distributed  in  the  North  At- 
lantic between  30°N  and  60°N,  the  South  Atlantic  south  of 
30°S,  and  also  in  the  eastern  South  Pacific  south  of  30°S. 

Nansenia  antarctica  new  species 

Figures  7,  8,  21 

HOLOTYPE.  LACM  10875  (one  of  three),  198.9  mm  SL, 
R/V  Eltanin  sta.  1204,  55°57'S,  159°23'W,  0560-1035,  10 
Aug.  1964,  I KMT,  0-4145  m. 

PARATYPES.  LACM  10875  (two  of  three)  (2,  197.8- 
207.5),  collection  data  as  for  holotype.  LACM  10658-15  (2, 
79.2-225.0),  R/V  Eltanin  sta.  858,  64°42'S,  78°34'W,  1905- 


% 


o * > 1 • — 

0 50  100  150  200 

S L (mm) 

Figure  8.  Nansenia  antarctica.  Predorsal,  head  lengths,  and  eye 
diameter  in  relation  to  standard  length. 


0030,  17-18  Nov.  1963,  I KMT;  USNM  247255  (1,  158.6), 
R/V  Eltanin  cr.  35,  sta.  2300,  52°00'S,  124°02'E,  20  Sep. 
1968,  I KMT,  0-750  m (bottom  depth  3914-4362  m). 

OTHER  MATERIALS.  LACM  11194  (2,  98.9-108.9), 
R/V  Eltanin  sta.  1661,  61°30'S,  108°26'W,  0134-0430,  26 
Apr.  1966,  I KMT,  0-5036  m;  LACM  11212(1,  49.9),  R/V 
Eltanin  sta.  1686,  57°39'S,  115°12'W,  1255-1540,  6 May 
1966,  I KMT,  0-4286  m;  USNM  247254  (7,  31.0-35.0), 
R/V  Eltanin  cr.  21,  sta.  20G,  0-850  m;  ISH  688/71  (1,  112.5), 
FFS  Walther  Herwig  sta.  363-III/71,  40°18'S,  39°04'W,  8 
Mar.  1971,  0-800  m;  ISH  542/71  (1,  118.2),  FFS  Walther 
Herwig  sta.  354-11/71,  39°19'S,  48°09'W,  6 Mar.  1971, 0-ca. 
2000  m. 

DIAGNOSIS.  Differs  from  other  Nansenia  in  having  three 
branchiostegal  rays,  47-50  vertebrae,  and  35-41  gill  rakers. 

DESCRIPTION.  D 9-10;  A 9-10;  P 12-14;  V 1 1-12;  gill 
rakers  10-13  + 1 + 23-27,  total  35-41  with  35(2),  37(2), 
38(2),  39(2),  and  41(1);  branchiostegals  three;  vertebrae  49- 
50  with  49(3)  and  50(5).  Pyloric  caeca  seven  in  one  specimen; 
27  irregularly  arranged  conical  teeth  on  head  of  vomer  and 
about  75  compressed  teeth  on  dentaries  on  one  specimen  of 
207.5  mm  SL. 

Body  slender,  anterior  part  slightly  compressed  in  speci- 
mens larger  than  190  mm,  subcylindrical  in  smaller  speci- 
mens. Caudal  peduncle  more  compressed  than  anterior  part 
of  body.  Body  depth-width  ratio  1. 1-1.7.  Eye  large,  diameter 
2. 4-2. 9 times  in  head  length,  with  its  dorsal  margin  pro- 
truding slightly  above  margin  of  head.  Golden  pigment, 
roughly  crescent-shaped,  on  posterior  half  of  iris.  The  ratio 
of  width  of  the  posterior  part  of  iris  to  that  of  the  anterior 
part  is  between  two  and  four,  whereas  the  ratio  is  less  than 
two  in  other  species  having  three  branchiostegal  rays.  Apha- 
kic space  extremely  well  developed  in  front  of  lens  (some- 


8 Contributions  in  Science,  Number  352 


Kawaguchi  and  Butler:  Fishes  of  the  Genus  Nansenia 


CM 


CM 


0 1 23456789  10 


12  13  14  15 


Figure  9.  Nansenia  megalopa.  ISH  2013-71  (one  of  the  two  catalogued),  133.0  mm  SL,  holotype. 


times  lens  placed  anteriorly),  its  horizontal  space  usually  three 
to  four  times  as  wide  as  the  width  of  the  iris  at  the  ante- 
riormost  part,  in  specimens  larger  than  100  mm  SL.  This 
space  is  less  well  developed  in  smaller  juveniles. 

Upper  end  of  pectoral  fin  base  just  below  midpoint  between 
the  lateral  line  and  ventral  margin  of  body,  its  lower  end 
separated  from  the  ventral  margin  by  a distance  1 .8-3.0  times 
as  long  as  its  base.  Origin  of  dorsal  fin  base  in  front  of  the 
center  of  the  body.  Ventral  fin  base  just  below  the  posterior 
end  of  the  dorsal  fin  base.  Adipose  fin  base  above  the  pos- 
terior half  of  anal  fin  base. 

Skin  badly  abraded  in  most  specimens,  light  brown  in 
alcohol.  Dark  colored  scale  pockets,  which  are  extremely 
fragile,  a few  sometimes  remaining  along  lateral  line  and  the 
dorsal  and  ventral  sides  of  body.  Caudal  and  adipose  fin  bases 
usually  slightly  pigmented. 

SIZE.  The  largest  specimens  studied,  207.5  mm,  had  ma- 
ture ovaries  with  developing  eggs. 

DISTRIBUTION.  This  species  occurs  in  the  southern  part 
of  the  subantarctic  region  between  the  antarctic  and  sub- 
tropical convergences  of  the  Pacific  (Fig.  21).  Its  distribution 
probably  extends  northward  to  the  subtropical  convergence 
in  the  Atlantic  sector. 

ETYMOLOGY.  Named  for  its  region  of  occurrence. 

SPECIES  WITH  FOUR  BRANCHIOSTEGAL  RAYS 

Nansenia  megalopa  new  species 

Figures  9,  2 1 

Nansenia  sp.  2 (?),  Parin  and  Golovan,  1976:251-252,  fig. 

2,  04°10'N,  off  west  Africa,  0-510  m. 

HOLOTYPE.  ISH  2013  a/71,  133.0  mm,  FFS  Walther 


Herwig  sta.  463-III/71,  08°11'S,  14°12'W,  8 Apr.  1971,  0- 
640  m. 

PARATYPES.  ISH  1730/71  (1,  126.5),  FFS  Walther  Her- 
wig sta.  447-III/71,  18°36'S,  04°18'W,  4 Apr.  1971;  ISH 
20 1 3b/7 1 (1,  108.5),  FFS  Walther  Herwig  sta.  463-III/71, 
08°1  l'S,  14°12'W,  8 Apr.  1971,  0-640  m;  ISH  2491/71  (1, 
139.5),  FFS  Walther  Herwig  sta.  486-III/71,  07°32'N, 
20°54'W,  14  Apr.  1971, 0-ca.  1300  m;  UF  29912  (1,  126.0), 
R/V  Geronimo  cr.  2,  sta.  82,  03°28'S,  00°14'W,  Gulf  of 
Guinea,  6 Aug.  1963,  0-710  m. 

DIAGNOSIS.  Differs  from  other  Nansenia  in  having  four 
branchiostegal  rays,  44-45  vertebrae  and  21-23  gill  rakers 
on  the  first  arch.  This  species  is  also  separable  from  all  other 
Nansenia  by  having  a very  large  eye,  eye  diameter  to  snout 
length  ratio  >2.8  to  1. 

DESCRIPTION.  D 9-10;  A 8-10;  P 12-13;  V 9-10;  gill 
rakers  6-7  + 1 + 14-15,  total  21-23  with  21(1),  22(2),  and 
23(2);  branchiostegal  rays  four;  vertebrae  44-45  with  44(3) 
and  45(2). 

Pyloric  caeca  not  counted;  teeth  on  head  of  vomer,  usually 
embedded  in  the  tooth  ridge  and  often  indiscernible;  35-50 
teeth  on  both  dentaries. 

Body  slender  and  subcylindrical,  body  depth  to  width  ratio 
1.2-1. 4 in  specimens  of  108.5-139.5  mm  SL.  Eye  large,  its 
diameter  2. 1 to  2.4  in  head  length.  A roughly  crescent-shaped, 
silvery  pigment  patch  on  the  posterior  half  of  the  iris;  pupil 
elliptical  with  a large  aphakic  space  in  front  of  the  lens.  Upper 
end  of  pectoral  fin  base  around  midpoint  between  lateral  line 
and  ventral  margin  of  body.  Dorsal  fin  base  originating  in 
front  of  center  of  body.  Ventral  fin  base  below  posterior  end 
of  dorsal  fin  base.  Adipose  fin  origin  above  posterior  third 
of  anal  fin  base.  Bases  of  ventral  and  adipose  fins  densely 
pigmented. 


Contributions  in  Science,  Number  352 


Kawaguchi  and  Butler:  Fishes  of  the  Genus  Nansenia  9 


H 10  mm 


Figure  10.  Nansenia  longicauda.  a.  SIO  76-7-9,  133  mm  SL,  holotype;  b.  ISH  60-66,  78.0  mm  SL,  paratype. 


SIZE.  A specimen  of  126.5  mm  SL  has  ripe  ovarian  eggs. 

DISTRIBUTION.  All  specimens  were  collected  in  the 
oceanic  area  of  the  tropical  Atlantic  between  10°N  and  20°S. 

ETYMOLOGY.  Nansenia  megalopa  from  Greek  megale, 
great,  and  from  ops,  eye,  referring  to  the  extraordinarily  large 
eye. 


Nansenia  longicauda  new  species 

Figures  10,  1 1,  22 

HOLOTYPE.  SIO  76-9,  133.0  mm,  30°37.0'N, 

147°24.0'W,  2027-2350,  10  Nov.  1971,IKMWT,  1800mwo. 

PARATYPES.  Pacific:  USNM  207530  (1,  104.6),  Univ. 
of  Hawaii,  samp.  no.  71-2-11,  21°20'N,  158°20'W,  28  Feb. 
1971,  610-650  m;  USNM  215703  (1,  97.2),  R/V  Townsend 
Cromwell,  CR-52,  sta.  16,  21°32.7'N,  158°21.8'W,  0355- 
1100,  12  Feb.  1971.  Atlantic:  ISH  60/66(1,78.0),  FFS  Wal- 
ther  Herwig  sta.  177-66,  33°45'N,  16°00'W,  10  May  1966, 
MT1600,  0-600  m. 

OTHER  SPECIMENS  EXAMINED.  Pacific  specimens: 
SIO  71-300  (1,  65.0),  R/V  Thomas  Washington  sta.  Aries 


9-H7,  27°24.5'N,  155°25.5'W,  0720-1100,  30  Sep.  1971, 
lOft.-IKMT,  3000  mwo;  USNM  207531  (1,  46.5),  Univ. 
of  Hawaii,  samp.  no.  71-6-10,  21°00'N,  158°20'W,  10  Jun. 
1971,  560-600  m;  USNM  215470  (1,  67.0),  R/V  Townsend 
Cromwell,  samp.  no.  73-8-29,  21°20'N,  158°20'W,  0745- 


% 

50  - 

40  - 


-o- 


o 

-o-® 

O O 


-®-  Pre  D 


30 

= 20 
“ 10 


HL 


t 0 


30  40  50  60  70  80  90  100  110  120  130  140 

S L ( mm  ) 


Figure  11.  Nansenia  longicauda.  Predorsal,  head  lengths,  and  eye 
diameter  in  relation  to  standard  length. 


10  Contributions  in  Science,  Number  352 


Kawaguchi  and  Butler:  Fishes  of  the  Genus  Nansenia 


Figure  12.  Nansenia  tenera.  a.  ZMUC  P 1 962 1-23  (one  of  the  three  cataloged),  129.0  mm  SL,  holotype;  b.  ISH  540-73,  117.0  mm  SL, 
paratype. 


1054,  29  Aug.  1973,  400-1 100  m;  SIO  71-302  (1,  67.0),  R/V 
Thomas  Washington  Aries  9-H9,  27°24.5'N,  155°25.5'E,  30 
Sep.  1971.  Atlantic  specimens:  USNM  246799  (1 , 41.5),  Acre 
6-22;  USNM  246800  (1,  39.1),  Acre  9-24.  Both  specimens 
were  collected  around  38°18'N  and  64°12'W. 

DIAGNOSIS.  Differs  from  other  Nansenia  in  having  four 
branchiostegal  rays,  23-27  gill  rakers  on  the  first  arch,  47- 
50  vertebrae  and  a slender  caudal  peduncle,  the  least  depth 
of  which  is  4. 3-5. 6 percent  of  SL.  The  body  shape  of  this 
species  resembles  that  of  N.  megalopa.  Higher  gill  raker  counts 
(23-27  vs.  21-23),  higher  vertebral  counts  (47-50  vs.  44- 
45)  and  longer  snout  (2.3-4. 1 vs.  2. 3-2.9  percent  of  SL) 
separate  N.  longicauda  from  N.  megalopa. 

DESCRIPTION.  D 10  (rarely  11);  A 10-11  (rarely  9);  P 
13-14  (rarely  1 1);  V 10-11  (rarely  9);  gill  rakers  7-8  + 1 + 
15-18,  total  23-27  with  23(1),  24(4),  25(2),  and  26(2);  bran- 
chiostegal rays  four;  vertebrae  47-50  with  47(1),  48(  1 ),  49(5), 
and  50(4).  These  counts  are  based  mainly  on  1 1 specimens, 
39.1-133.0  mm  SL.  Pyloric  caeca  not  counted.  Irregularly 
sized  and  arranged  conical  teeth  on  head  of  vomer,  which 
are  embedded  deeply  in  tooth  ridge,  about  24;  compressed 
teeth  closely  set  on  dentaries,  about  50  in  a 104.6-mm  spec- 
imen. 

Body  slender  and  subcylindrical  with  body  depth  to  width 
ratio  1.3-1. 6 in  specimens  larger  than  45  mm  SL.  The  ratio 
is  larger  in  smaller  specimens.  Eye  very  large,  indicating  con- 
siderable allometric  growth  (Fig.  11);  its  diameter  varying 
linearly  from  6.4  percent  of  SL  in  a 39.1 -mm  specimen  to 
8.8  percent  in  a 133.0-mm  specimen;  eye  diameter  2. 2-2. 6 


times  in  head  length  in  specimens  larger  than  50  mm,  and 
2. 7-3.0  times  in  a specimen  smaller  than  50  mm. 

Dorsal  margin  of  eye  protuding  above  dorsal  margin  of 
head  in  a specimen  larger  than  100  mm  SL.  Pupil  oval,  with 
aphakic  space  developed  anteriorly.  Base  of  uppermost  pec- 
toral fin  at  or  slightly  higher  than  midpoint  between  lateral 
line  and  ventral  margin  of  the  body.  Origin  of  dorsal  fin  base 
well  behind  center  of  body.  Predorsal  length  less  than  47 
percent  of  standard  length  with  smaller  values  in  larger  spec- 
imen due  to  allometric  growth  (Fig.  1 1).  Base  of  outermost 
ventral  fin  below  or  just  behind  posterior  end  of  dorsal  fin 
base.  Adipose  fin  base  just  in  front  of  vertical  through  center 
of  anal  fin  base. 

Tip  of  snout,  upper  jaw,  anterior  part  of  gular  area,  bases 
of  ventral  and  caudal  fins  densely  pigmented. 

SIZE.  The  largest  specimen  is  the  133.0-mm  SL  holotype. 

DISTRIBUTION.  Nansenia  longicauda  occurs  in  the  sub- 
tropical and  temperate  waters  of  the  Pacific  and  Atlantic 
oceans.  Based  on  collection  data,  this  species  appears  to  be 
an  oceanic  mesopelagic  species  (Fig.  22). 

ETYMOLOGY.  The  specific  name  is  derived  from  Latin, 
longus  meaning  long  and  cauda  meaning  tail  referring  to  the 
long  and  slender  caudal  peduncle. 

Nansenia  tenera  new  species 

Figures  12,  21 

HOLOTYPE.  ZMC  PI 962 1-23  (1,  129.0),  Dana  sta. 
3975-1,  35°42'S,  18°37'E,  31  Jan.  1930,  3000  mwo. 


Contributions  in  Science,  Number  352 


Kawaguchi  and  Butler:  Fishes  of  the  Genus  Nansenia  1 1 


-I  10  mm 


-i  10  mm 


C 


i 1 10  mm 


Figure  13.  Nansenia  ardesiaca.  a.  FMNH  57086,  164.5  mm  SL,  holotype  from  off  Japan;  b.  USNM  93354,  136.0  mm  SL,  one  of  the  seven 
paratypes  of  N.  schmitti  from  off  the  Philippines;  c.  ZMUC  P19624-25,  183.5  mm  SL,  from  off  the  northeast  coast  of  South  Africa. 


PARATYPES.  ZMC  PI 962 1-23  (2,  1 12.0-1  17.4),  collec- 
tion data  as  for  the  holotype;  ZMC  not  catalogued  (1,  105.6), 
Dana  sta.  3975-11,  35°42'S,  18°37'E,  31  Jan.  1930,  2500  mwo; 
ISH  1009/73  (1,  117.0),  FFS  Wa/ther  Herwig  696-73, 
55°39'N,  25°42'W,  22  Sep.  1973,  400-410  m;  ISH  829/71 
(1,  101.0),  FFS  Walther  Herwig  376/71,  39°55'S,  26°02'W, 
11  Mar.  1971,  0-2000  m. 

DIAGNOSIS.  Differs  from  other  species  of  Nansenia  in 
having  four  branchiostegal  rays,  42-43  vertebrae,  and  43- 
46  gill  rakers. 


DESCRIPTION.  D 1 1-12;  A 8-9;  P 1 1-12;  V 1 1-12;  gill 
rakers  13-15  + 1 + 28-32,  total  43-46  with  43(1),  44(3), 
45(1),  and  46(1);  branchiostegal  rays  four;  vertebrae  42  in 
one  North  Atlantic  specimen,  43  in  the  five  South  Atlantic 
specimens.  Pyloric  caeca  7-9  in  three  specimens;  about  16 
conical  teeth  irregularly  arranged  on  head  of  vomer;  about 
60  compressed  teeth  on  dentaries  in  a 1 12-mm  specimen. 

Eye  large,  its  diameter  2. 4-2. 7 in  head  length;  nearly  one- 
third  to  one-quarter  of  eye  protruding  above  the  dorsal  mar- 
gin of  head.  A golden  colored,  crescent-shaped  pigment  on 


12  Contributions  in  Science,  Number  352 


Kawaguchi  and  Butler:  Fishes  of  the  Genus  Nansenia 


1 0 mm 


Figure  14.  Nansenia  tenuicauda.  ISH  1010-71,  69.0  mm  SL,  paratype. 


posterior  half  of  iris.  Aphakic  space  developed;  pupil  acorn- 
shaped  with  tip  anteriorly  directed.  This  type  of  pupil  was 
not  observed  in  any  other  species. 

Upper  end  of  pectoral  fin  base  below  midpoint  between 
lateral  line  and  ventral  margin  of  body;  its  lower  end  above 
ventral  margin  of  body  by  a space  greater  than  length  of 
pectoral  fin  base.  Dorsal  fin  originating  in  front  of  center  of 
body.  Ventral  fin  base  above  posterior  half  of  anal  fin  base 
in  the  South  Atlantic  specimens,  but  above  the  midpoint  in 
one  specimen  from  the  North  Atlantic. 

Body  color  of  preserved  specimens  light  brown.  The  entire 
body  surface  of  three  specimens,  especially  the  ventral  side, 
is  sparsely  covered  with  guanine,  which  resembles  silver  pow- 
der. Caudal  fin  base  densely  pigmented  in  the  specimens  from 
the  South  Atlantic,  but  the  pigmentation  extends  forward 
becoming  sparser  anteriorly  in  the  North  Atlantic  specimen. 
SIZE.  A specimen  of  1 12.0  mm  SL  has  maturing  gonads. 
DISTRIBUTION.  Five  specimens  were  collected  in  the 
South  Atlantic  between  35°S  and  39°S,  and  one  specimen 
from  the  subarctic  region  of  the  North  Atlantic  (55°39'N). 

ETYMOLOGY.  The  specific  name  tenera  is  from  Latin, 
soft,  referring  to  the  consistency  of  the  body. 

Nansenia  ardesiaca  Jordan  and  Thompson,  1914 

Figures  13,  22 

Nansenia  groenlandica:  Tanaka,  1911:15-16,  pi.  Ill,  fig.  13 
(non  Reinhardt,  1840)  (Sagami  Bay,  Japan). 

Nansenia  ardesiaca  Jordan  and  Thompson,  1914:210-21 1, 
pi.  XXIV,  fig.  2 (orig.  descr.,  off  Japan). 

Nansenia  tanakai  Schmidt,  1918:15  (orig.  descr.  Japan). 
Bathymacrops  microlepis  Gilchrist,  1922:53-54,  pi.  IX,  fig. 

2 (orig.  descr.,  east  coast  of  South  Africa). 

Microstoma  (Euproserpa)  schmitti  Fowler,  1934:256-257,  fig. 
18  (orig.  descr.  Philippines). 

Nansenia  macrolepis:  Cohen,  1958:56  (east  coast  of  South 
Africa). 


Nansenia  robusta  Abe,  1976:27-31,  figs.  1-6  (orig.  descr. 

South  China  Sea). 

MATERIALS.  Off  Japan:  FMNH  57086,  164.5  mm  SL, 
Okinose,  Sagami  Bay,  central  Japan  (holotype  of  N.  ardesia- 
ca)', KYO  4434  (2,  157.5,  144.0),  southern  Japan,  KU  7013 
(1,1 20.0),  Mimase  fish  market,  Shikoku  Is.,  Japan;  KU  13012 
(1,  141.5),  Tosa  Bay,  Shikoku  Is.,  430-460  m;  KU  131 25— 
13130,  13243,  13244,  13279  (9,  124.0-155.0),  15  km  east 
of  Ashizuri  Misaki  Pt.,  Shikoku  Is.,  bottom  trawl  420- 
555  m. 

Southeast  Asian  Seas:  USNM  92327  (holotype  of  N. 
schmitti)  (1,  194.5),  R/V  Albatross,  sta.  D5445,  Atalaya  Pt., 
Batag  Is.,  off  Samar,  the  Philippines,  3 Jun.  1909;  USNM 
93354  (1,  136.0),  R/V  Albatross,  sta.  5589,  04°12.10'N, 
118°38.08'E,  Mabul  Is.,  off  the  Philippines  (one  of  seven 
paratypes  of  N.  schmitti)',  KU  15792  (1,  88.5,  damaged), 
R/V  Hakuho  Maru,  Cr.  KH72-1,  sta.  20,  05°40.9'N, 
1 19°46.3'E,  bottom  trawl  460-500  m;  KU  16644  (1,  207.0), 
R/V  Hakuho  Maru,  KH72-1,  sta.  26,  09°27.0'S,  127°58.6'E, 
Timor  Sea,  bottom  trawl  690-850  m. 

Off  South  Africa:  USNM  2034391-10  (10,  174.0-192.0), 
R/V  Anton  Bruun,  22°25'S,  35°54'E,  shrimp  trawl,  0-740  m; 
ZMC  P19624-25  (2,  183.5,  185.0),  Galathea  exped.  1950- 
1952,  sta.  203,  25°36'S,  35°21'E  off  Natal,  2015  m;  USNM 
not  catalogued,  SOSC  ref.  no.  1701-2  (2,  150.0,  165.0),  IIO 
Exped.  R/V  Anton  Bruun,  Cr.  no.  8,  sta.  397C,  26°07'S, 
34°1  l'E,  bottom  trawl  600-665  m;  CAS-SU-31501  (1,  127.5), 
off  South  Africa. 

DIAGNOSIS.  Differs  in  having  four  branchiostegal  rays, 
predorsal  length  less  than  49  percent  of  SL,  length  of  caudal 
peduncle  not  more  than  1 5 percent  of  SL,  number  of  gill 
rakers  on  first  arch  27-35. 

DESCRIPTION.  D 9-10;  A 9-10;  P 12-14  (rarely  1 1);  V 
11  (rarely  10,  12);  gill  rakers  9-11  + 1 + 19-23,  total  29- 
35  with  29(1),  30(1),  31(1),  32(5),  34(2),  and  35(2)  in  12 
specimens  from  off  Japan;  9 + 1 + 18-21,  total  27-31  with 


Contributions  in  Science,  Number  352 


Kawaguchi  and  Butler:  Fishes  of  the  Genus  Nansenia  13 


Figure  15.  Nansenia  pelagica.  ISH  649-74,  102.0  mm  SL,  holotype. 


27(1),  30(1),  and  31(1)  in  three  specimens  from  off  the  Phil- 
ippines; 8-10  + 1 + 19-21,  total  28-31  with  28(1),  29(2), 
30(7),  and  3 1(4)  in  14  specimens  from  off  South  Africa;  bran- 
chiostegal  rays  four;  vertebrae  46-48  with  46(6),  47(12),  and 
48(2). 

Gill  raker  counts  were  rather  variable  and  higher  on  the 
average  in  the  specimens  from  off  Japan.  Ranges  of  these 
counts,  however,  overlap  among  specimens  from  the  three 
different  regions.  No  significant  geographical  variation  was 
found  in  other  counts  and  measurements. 

Pyloric  caeca  8,  8,  and  9 in  three  specimens  from  off  South 
Africa,  and  7,  9,  and  9 in  two  Japanese  specimens;  conical 
teeth  on  vomerine  series  25-35  and  teeth  on  dentaries  65- 
75. 

Eye  large,  diameter  2. 3-2. 5 times  in  head  length,  dorsal 
margin  protruding  above  the  margin  of  head.  A roughly  cres- 
cent-shaped, yellowish  pigment  patch  present  in  the  posterior 
half  of  iris.  Aphakic  space  well  developed.  Anterior  hori- 
zontal space  nearly  equal  to  width  of  iris,  about  twice  width 
of  posterior  space.  Aphakic  space  probably  less  well  devel- 
oped in  smaller  specimens,  as  is  the  case  in  other  species. 

Upper  end  of  pectoral  fin  base  above  midpoint  between 
lateral  line  and  ventral  margin  of  body,  its  lower  end  well 
above  ventral  margin  of  body  by  a distance  more  than  (usu- 
ally 1.5  to  twice)  length  of  pectoral  fin  base.  Dorsal  fin  origin 
in  front  of  center  of  body.  Ventral  fin  base  below  or  just 
behind  posterior  end  of  dorsal  fin  base.  Adipose  fin  base 
above  posterior  half  of  anal  fin  base.  Tip  of  mouth,  snout, 
adipose,  caudal  and  ventral  fin  bases  pigmented. 

SIZE.  This  species  reaches  about  200  mm  SL.  A specimen 
of  196  mm  SL  is  reported  to  have  ripe  eggs  (Abe,  1976). 

DISTRIBUTION.  Based  on  previous  records,  N.  ardesia- 
ca  occurs  in  slope  waters  of  Southeast  Asian  Seas,  as  far  north 


as  off  southern  Japan.  It  also  occurs  in  slope  waters  off  the 
east  coast  of  South  Africa  (Fig.  22). 

Nansenia  tenuicauda  new  species 

Figures  14,  22 

HOLOTYPE.  ISH  658/71, 84.0  mm,  FFS  Walther  Herwig 
sta.  363-11/71,  40°18'S,  39°12'W,  8 Mar.  1971,  MT1600,  0- 
328  m. 

PARATYPES.  ISH  1010-71  (2,  70.5,  69.0),  FFS  Walther 
Herwig  sta.  399-11/71,  40°34'S,  09°50'W,  18  Mar.  1971, 
MT1600,  0-310  m. 

DIAGNOSIS.  Differs  in  having  four  branchiostegal  rays, 
predorsal  length  less  than  49  percent  of  SL  and  38-42  gill 
rakers  on  the  first  arch.  Slender  caudal  peduncle  the  least 
depth  of  which  is  less  than  5.5  percent  of  SL  also  distinguish 
this  species  from  other  species  of  Nansenia. 

DESCRIPTION.  D 10-11;  A 8-10;  P 11-13;  V 11-12; 
gill  rakers  1 1-1 5 + 1 + 25-26,  total  38-42  with  38(1),  39(1), 
and  42(1);  branchiostegal  rays  four;  vertebrae  46.  Counts  are 
based  on  three  specimens. 

Pyloric  caeca  eight  in  one  specimen.  Conical  teeth  ca.  eight, 
irregularly  arranged  on  head  of  vomer  and  ca.  26  compressed 
teeth  on  dentaries.  A pair  of  teeth  on  the  symphysis  longest 
of  all.  Other  teeth  of  lower  jaw,  as  usually  observed  in  other 
species  of  Nansenia.  are  progressively  smaller  from  the  back 
of  each  dentary  to  the  front. 

Body  subcylindrical,  with  a depth  to  width  ratio  of  1.3- 
1 .4.  Eye  large,  its  diameter  2. 8-3.0  times  in  head  length;  pupil 
ovoid  in  shape  with  aphakic  space  developed  posteriorly. 

Upper  end  of  pectoral  fin  base  just  below  the  midpoint 
between  lateral  line  and  ventral  margin  of  the  body.  Dorsal 
fin  origin  well  in  front  of  center  of  the  body.  Ventral  fin  base 


14  Contributions  in  Science,  Number  352 


Kawaguchi  and  Butler:  Fishes  of  the  Genus  Nansenia 


below  the  posterior  end  of  dorsal  fin  base.  Adipose  fin  base 
above  base  of  the  last  two  anal  fin  rays. 

Base  of  caudal  fin  densely  pigmented,  but  pigment  becom- 
ing progressively  sparser  anteriorly.  Body  color  of  preserved 
specimens  light  brown. 

SIZE.  The  specimens  examined,  69.0-84.0  mm  SL,  are 
probably  juveniles. 

DISTRIBUTION.  Restricted  to  the  area  near  the  sub- 
tropical convergence  in  the  South  Atlantic,  around  40°S. 

ETYMOLOGY.  From  Latin,  tenuis,  slender,  and  from 
cauda,  tail,  in  reference  to  the  slender  caudal  peduncle. 

Nansenia  pelagica  new  species 

Figures  15,  16,  22 

HOLOTYPE.  ISH  649/74  (1,  102.0),  sta.  A.D.  61-11/74, 
02°27'N,  34°52'W,  24  Jan.  1974,  MT1600,  0-350  m. 

PARA  TYPES.  Atlantic:  ISH  1865/66  (1,75.5),  FFS  Wal- 
ther  Herwig  sta.  182-66,  10°46'N,  23°54'W,  16  May  1966, 
0-300  m;  ISH  442/66  (1,  93.0),  FFS  Walther  Herwig  sta. 
1 84-66,  06°25'N,24°34'W,  17May  1 966,  0-320  m;  ISH  57 1/ 
66  (1,  101.0),  FFS  Walther  Herwig  sta.  186-66,  01°24'S, 
25°58'W,  19  May  1966,  0-330  m;  ISH  619/66  (1, 64.8),  FFS 
Walther  Herwig  sta.  187-66,  05°34'S,  26°58'W,  20  May  1966, 
0-320  m;  ISH  2066/71  (2,  101.5,  107.5),  FFS  Walther  Her- 
wig 467-71,  05°30'S,  16°28'W,  9 Apr.  1971,  0-1900  m. 
Pacific:  USNM  207532  (1,  83.7),  21°20'N,  158°20'W  (off 
Hawaii),  16  Sep.  1970,  0-725  m. 

OTHER  MATERIALS.  Atlantic:  ISH  928/68  (1,  58.5), 
FFS  Walther  Herwig  sta.  17-68,  04°43'S,  26°39'W,  MT1600, 
0-ca.  2000  m.  Pacific:  USNM  201704  (1,  27.0),  R/V  Swan, 
28°48'N,  1 4 1 °59'W,  6 ft.  - 1 KMT,  200-247  m;  USNM  212102 
(2,  ca.  60.5,  ca.  54,  damaged),  from  the  stomach  of  a Thunnus 
albacares  collected  off  the  New  Hebrides  Is.,  20°00'S, 
170°03'E. 

DIAGNOSIS.  Differs  from  other  Nansenia  in  having  four 
branchiostegal  rays  and  38-39  vertebrae.  Although  it  is  sim- 
ilar to  N.  atlantica,  and  their  distributions  overlap  in  the 
tropical  Atlantic,  N.  pelagica  differs  from  N.  atlantica  in 
having  lower  vertebral  counts  (38-39  vs.  41-42),  a longer 
head  (27-30  percent  of  SL  vs.  20-26  percent),  and  a dark 
pigment  spot  on  the  gular  area  that  becomes  obscure  with 
growth  in  specimens  larger  than  90  mm  SL. 

DESCRIPTION.  D 9-10;  A 8-9;  P 9-10  (rarely  1 1);  V 
10-1 1;  gill  rakers  10-13  + 1 + 24-27,  total  36-4 1 with  36(2), 
37(1),  38(3),  39(2),  and  41(1);  branchiostegal  rays  four,  ver- 
tebrae 38-39  with  38(6)  and  39(4).  Counts  are  based  on  10 
specimens,  58.5-107.5  mm.  Gill  raker  count  of  the  one  Pa- 
cific specimen  was  higher  than  counts  from  the  Atlantic  (41 
vs.  36-38).  Other  counts  showed  no  remarkable  difference 
between  the  Pacific  and  Atlantic  specimens. 

Pyloric  caeca  seven  and  eight  in  the  two  Atlantic  speci- 
mens. About  35  conical  teeth  on  vomerine  series  and  about 
60  teeth,  resembling  a closely  spaced  picket  fence,  on  the 
lower  jaw  of  the  specimens  of  68.4  mm  SL. 

Body  slender  and  slightly  compressed,  with  a body  depth- 
width  ratio  of  1.3-1. 6 in  specimens  larger  than  58  mm  SL. 
Eye  large,  diameter  2. 7-2. 8 in  head  length  in  specimens  58.5- 


% 

60 

O 

o 

50 

o 

o 

— o o__ 

Pre  D 

CO 

40- 

o 

30 

o 

H L 

= 

o ° 

o 

^ ° ~ 

20 

10 

E D 

o 

o 

0 . * • * • * • * • — 

10  20  30  40  50  60  70  80  90  100  110 

S L ( mm  ) 

Figure  16.  Nansenia  pelagica.  Predorsal,  head  lengths,  and  eye 
diameter  in  relationship  to  standard  length. 

93.0  mm  SL,  and  2.4  to  2.5  times  in  head  length  in  specimens 
99.5-107.5  mm  SL.  Yellowish  pigment,  in  a roughly  cres- 
cent-shaped patch,  develops  on  posterior  half  of  iris  with 
growth.  Aphakic  space  developed  in  front  of  lens.  This  space 
is  inconspicuous  in  juveniles. 

Upper  end  of  pectoral  fin  base  just  below  midpoint  between 
lateral  line  and  ventral  margin  of  body,  its  lower  end  above 
ventral  margin  of  body  by  a distance  about  1.5  times  as  long 
as  length  of  pectoral  fin  base.  Origin  of  dorsal  fin  base  just 
behind  center  of  body.  Adipose  fin  base  over  anterior  half 
of  anal  fin  base. 

Body  skinned  in  most  specimens,  but  lateral  line  pockets 
remaining  in  some.  Bases  of  caudal,  dorsal,  ventral  and  adi- 
pose fins,  and  dorsal  margin  darker  than  lateral  side  of  body. 
Inside  of  opercle  lined  with  dark  membrane.  Dark  pigment 
spot  on  gular  area  clearly  recognizable,  especially  in  juveniles 
less  than  70  mm  SL. 

SIZE.  The  largest  specimen  examined,  probably  an  adult, 
was  107.5  mm  SL. 

DISTRIBLITION.  Nansenia  pelagica  occurs  in  the  trop- 
ical Atlantic  between  1 5°N  and  1 0°S.  In  the  Pacific  it  is  known 
only  from  the  subtropical  eastern  North  Pacific  between  Ha- 
waii and  North  America  and  the  tropical  western  South  Pa- 
cific off  the  New  Hebrides  Islands. 

ETYMOLOGY.  In  reference  to  the  pelagic  life  of  this 
species. 

Nansenia  atlantica  Blache  and  Rossignol,  1962 

Figures  17,  21 

Nansenia  atlantica  Blache  and  Rossignol,  1 962: 105-106,  fig. 

1 (orig.  descr.  16  mm  SL,  01°55'S,  8°30'E;  15  mm  SL, 

03°36'S,  09°10'E). 

MATERIALS.  ISH  335/66  (2,  88.5,  one  damaged),  FFS 
Walther  Herwig  sta.  182-66,  10°46'N,  23°54'W,  0-300  m; 
ISH  457/68  (4,  90.5-1  12.8,  one  damaged),  FFS  Walther  Her- 
wig  sta.  11-1-68,  16°14'N,22°24'E,  0-160  m;  ISH  686/68  (1, 
ca.  161,  damaged),  FFS  Walther  Herwig  sta.  13-11-68, 
08°2 1 'N,  24°10'W,  0-520  m;  ISH  13-11/68  (1,  167.5),  FFS 
Walther  Herwig  sta.  1 1-III-68,  16°08'N,  22°22'W,  0-580  m. 


Contributions  in  Science,  Number  352 


Kawaguchi  and  Butler:  Fishes  of  the  Genus  Nansenia  15 


-i  1 0 mm 


H 


Figure  17.  Nansenia  atlantica.  ISH  475-68,  104.5  mm  SL,  showing  proximal  part  of  adipose  fin  densely  pigmented. 


DIAGNOSIS.  Differs  from  other  species  of  Nansenia  in 
having  four  branchiostegal  rays,  origin  of  dorsal  fin  base  in 
front  of  center  of  the  body,  41-42  vertebrae  and  dark  pigment 
spot  at  the  base  of  adipose  fin.  This  species  is  closely  related 
to  N.  oblita  as  all  counts  overlap.  The  densely  pigmented 
proximal  part  of  the  adipose  fin  base  in  N.  atlantica  distin- 
guishes the  two. 

DESCRIPTION.  D 9-10;  A 8-9;  P 12-13;  V 10-11;  gill 
raker  on  the  first  arch  9-13  + 1 + 19-23,  total  30-36  with 
30(1),  31(2),  33(2),  35(1),  and  36(1);  branchiostegal  rays  four; 


vertebrae  41-42  with  41(3)  and  42(5).  Counts  are  based  on 
five  to  eight  specimens. 

Pyloric  caeca  seven  in  three  specimens  and  nine  in  one 
specimen.  Conical  teeth  on  head  of  vomer  35,  about  140 
teeth  on  dentaries  in  one  specimen. 

Body  slender  and  compressed  laterally  with  body  depth- 
width  ratio,  1.4-1. 6.  Eye  diameter  2. 7-3.0  times  in  head 
length,  viewed  laterally  its  dorsal  margin  touching  or  pro- 
truding slightly  higher  than  dorsal  margin  of  head.  Yellowish 
iridescent  pigment  pattern,  roughly  crescent-shaped,  recog- 


i 1 10  mm 


Figure  18.  Nansenia  oblita.  USNM  92241,  39.1  mm  SL,  juvenile. 


16  Contributions  in  Science,  Number  352 


Kawaguchi  and  Butler:  Fishes  of  the  Genus  Nansenia 


10  mm 


Figure  19.  Nansenia  crassa.  LACM  9808-32,  165.2  mm  SL. 


nizable  in  the  posterior  half  of  iris.  Pupil  elliptical  with  aphakic 
space  in  front  of  lens. 

Upper  end  of  pectoral  hn  base  well  below  midpoint  be- 
tween lateral  line  and  ventral  margin  of  body.  Dorsal  hn  base 
originating  just  below  center  of  body.  Ventral  hn  base  just 
in  front  of  posterior  end  of  dorsal  hn  base.  Adipose  hn  base 
above  center  of  anal  hn  base. 

Body  skin  brown  and  easily  rubbed  off.  The  bases  of  adi- 
pose and  caudal  hns  densely  pigmented. 

REMARKS.  The  original  description  of  N.  atlantica  was 
based  on  two  juvenile  specimens  of  16  and  1 5 mm  SL.  There 
is  a considerable  size  gap  between  the  type  material  and  the 
smallest  specimen  available.  Therefore,  the  present  speci- 
mens are  assigned  to  N.  atlantica  based  on  the  coincidence 
of:  (1)  counts  of  branchiostegal  and  anal  hn  rays;  (2)  locality; 
(3)  a wholly  pigmented  dark  body  surface.  In  the  tropical 
Atlantic,  two  species  of  the  Br-4  group  occur,  namely  N. 
atlantica  and  N.  pelagica.  Juveniles  of  the  latter  differ  from 
the  former  in  the  absence  of  complete  pigmentation  on  the 
body. 

SIZE.  The  largest  specimens  of  167.5  and  161  mm  SL, 
collected  in  January,  were  mature  males.  An  immature  fe- 
male of  1 12.0  mm  SL  was  collected  at  the  same  time. 

DISTRIBUTION.  All  specimens,  including  the  types,  were 
collected  in  the  eastern  tropical  Atlantic  between  20°N  and 
10°S  (Fig.  21). 

Nansenia  oblita  (Facciola,  1887) 

Figures  18,  21 

Microstoma  argent eum  oblitum  Facciola,  1887:193  (orig. 

descr.,  Messina,  Mediterranean). 

Microstoma  rissoanum  Sarato,  1890  (orig.  descr.,  Nice). 
Microstoma  oblitum:  Belloti,  1888:224,  hgs.  3a,  3aA. 
Nansenia  oblita:  Schmidt,  1918:19-22,  hgs.  13-14;  Cohen, 

1958:55. 


Doubtful  references: 

Leuroglossus  oblitus:  Kuroda,  1947:25  (listed,  no  descrip- 
tion, material  unavailable);  Kuroda,  1951:318. 

Nansenia  oblita:  Matsubara,  1955:216  (cited  from  Kuroda, 
1947). 

MATERIALS.  USNM  40075  (2,  40.5-48.5),  from  Mes- 
sina, originally  deposited  in  Royal  Zool.  Mus.  Florence; 
USNM  92241  (3,  34.2-46.0),  Mediterranean,  originally  de- 
posited in  Milano  Mus. 

DIAGNOSIS.  Differs  in  having  four  branchiostegal  rays, 
predorsal  length  longer  than  50  percent  of  standard  length, 
gill  raker  count  28-30  and  proximal  part  of  adipose  hn  un- 
pigmented. 

DESCRIPTION.  D 10-11;  A 9-10;  P 10-11;  V 10-11; 
gill  rakers  on  the  hrst  arch  7-8  + 1 + 20-21,  total  28-30 
with  28(1)  and  30(2);  branchiostegal  rays  four  on  three  spec- 


°/o 

70 


0 1 1 1 t t— 

0 50  100  150  200  230 

S L ( mm  ) 


Figure  20.  Nansenia  crassa.  Predorsal,  head  lengths,  and  eye  di- 
ameter in  relation  to  standard  length. 


Contributions  in  Science,  Number  352 


Kawaguchi  and  Butler:  Fishes  of  the  Genus  Nansenia  17 


Figure  21.  Distributions  of  specimens  examined  of  seven  species  of  Nansenia.  A single  symbol  may  indicate  more  than  one  record. 


imens;  vertebrae  42-45  with  42(1),  43(1),  44(2),  and  45(1). 
Fin  ray  counts  are  based  on  four  specimens. 

Pyloric  caeca  eight  in  one  specimen  (seven  reported  by 
Schmidt,  1918);  about  18  conical  teeth  irregularly  arranged 
on  head  of  vomer  and  about  30  compressed  teeth  on  den- 
taries  in  a specimen  of  48.5  mm  SL.  Measurements  are  based 
on  four  juveniles,  34.2-48.5  mm.  The  present  counts  and 
measurements  are  not  very  different  from  those  presented 
for  Mediterranean  specimens  by  Schmidt  (1918),  except  for 
the  slightly  lower  pectoral  and  ventral  fin  ray  counts  in  our 
material. 

Body  laterally  compressed  and  stubby  rather  than  slender 
with  body  depth-width  ratio,  1.3-1. 6.  Eye  diameter  3.0  to 
3.4  in  head  length  in  juveniles  (this  ratio  may  be  higher  in 
adults  due  to  negative  allometry  as  observed  in  the  closely 
related  N.  crassa)\  pupil  nearly  round  in  a specimen  of  34.2 
mm  SL,  horizontally  elliptical  in  specimens  larger  than  39. 1 
mm. 

Upper  end  of  pectoral  fin  base  above  or  on  midpoint  be- 
tween lateral  line  and  ventral  margin  of  body,  its  lower  end 
separated  from  ventral  margin  of  body  by  a distance  more 


than  length  of  pectoral  fin  base.  Dorsal  fin  origin  behind 
center  of  body.  Ventral  fin  base  below  posterior  end  of  dorsal 
fin  base.  Adipose  fin  base  above  the  middle  of  anal  fin  base 
or  slightly  in  front  of  it. 

Entire  body  covered  with  guanine;  base  of  caudal  and  pro- 
current caudal  fin  rays  pigmented. 

SIZE.  Largest  specimen  reported  by  Belloti  (1888)  about 
1 8 cm.  Therefore  the  present  material  is  probably  all  juvenile. 

DISTRIBUTION.  Common  in  the  western  Mediterra- 
nean (Schmidt,  1918).  Although  Schmidt  (1918)  reported 
two  juveniles,  21  and  31.5  mm,  from  the  temperate  eastern 
Atlantic  south  of  the  British  Isles  (48°43'N,  12°05'W),  we 
have  seen  only  Mediterranean  specimens,  in  spite  of  consid- 
erable fishing  effort  by  FFS  Walther  Herwig  in  the  eastern 
North  Atlantic. 

Nansenia  crassa  Lavenberg,  1965 

Figures  19,  20,  21 

Nansenia  crassa  Lavenberg,  1965:282-285,  fig.  1,  table  1 
(orig.  descr.). 


18  Contributions  in  Science,  Number  352 


Kawaguchi  and  Butler:  Fishes  of  the  Genus  Nansenia 


MATERIALS.  LACM  4425,  holotype,  212.0  mm,  R/V 
Ve/ero,  sta.  IV8296,  33°1 7'N,  118°40'W,  1 336-1642,  9 Nov. 
1962,  0-580  m;  LACM  4426,  paratype,  202  mm,  R/V  Ve- 
lero,  sta.  IV7374,  33°28'N,  1 18°18'W,  0200-0550,  29  Jim., 
1961,  0-865  m;  LACM  4427,  paratype,  266  mm  (275  mm 
in  original),  R/V  Velero,  sta.  IV8934,  33°14'N,  118°33'W, 
1559-1858,  18  Sep.  1963,  0-485  m;  LACM  9031-6(1,26.3), 
R/V  Velero IV,  33°20.0'N,  1 1 8°38.22' W, 0704-1022,  10ft.- 
IKMT,  0-504  m;  LACM  9068-1  (1,  44.5),  R/V  Velero  IV, 
33°36'48"N,  118°26'26"W,  1241-1445,  lOft.-IKMT,  0-ca. 
500  m;  LACM  9681  (2,  33.5,  39.0),  R/V  Velero,  sta.  11616, 
Valero  Basin,  Mexico;  LACM  9808-32  (1,  165.2),  R/V  Ve- 
lero, San  Clemente  Basin,  Calif.;  LACM  9809-1  (2,  34.0, 
35.0),  R/V  Velero,  sta.  10682,  29°17'00"N,  118°10'30"W,  10 
ft.-IKMT;  LACM  9965-9  (4,  19.0-22.0),  R/V  Velero,  sta. 
9891,  San  Juan  Seamount;  LACM  35549  (1,  203.6),  R/V 
Velero,  San  Clemente,  Calif.;  SIO  H51-190  (1,  238.0), 
32°49'N,  1 17°41'W,  0-548  m;  SIO  57-87  (1,  93.0),  29°I5'N, 
! 26°07  W.  off  Baja  Calif. 

DIAGNOSIS.  Differs  from  other  species  of  the  genus  in 
the  following  characters:  ( 1 ) four  branchiostegal  rays;  (2)  pre- 
dorsal length  more  than  49  percent  of  standard  length;  (3) 
43-46  (usually  44-45)  vertebrae;  (4)  length  of  caudal  pe- 
duncle more  than  13  percent  of  SL  and  depth  of  caudal 
peduncle  less  than  8.0  percent  of  SL  (CPL/CPD  ratio  1.7- 
2.2)  in  the  specimens  more  than  30  mm  SL.  Gill  raker  count 
(35-37)  also  distinguishes  this  species  from  the  closely  related 
N.  oblita  (28-30). 

DESCRIPTION.  D 9-10;  A 9 (rarely  8);  P 11-13;  V 10- 
1 1;  gill  rakers  on  the  first  arch  12-14  + 1 + 22-23,  total  35- 
37  with  35(2),  36(2),  and  37(2);  branchiostegal  rays  four; 
vertebrae  43-46  with  43(1),  44(3),  45(5),  and  46(2). 

Pyloric  caeca  seven  in  two  specimens;  conical  teeth  in 
vomerine  series  20-35,  teeth  on  dentaries  150-180  in  five 
specimens  larger  than  160  mm  SL. 

Morphometries  based  on  five  juveniles  and  seven  adults 
show  remarkable  allometric  growth  in  head  and  predorsal 
lengths  (Fig.  20). 

Eye  diameter  3.3  to  4.0  in  head  length  with  larger  values 
in  smaller  specimens  due  to  allometry;  viewed  laterally  dorsal 
margin  (supraorbital  bone)  protruding  slightly  above  dorsal 
margin  of  head  (less  in  juveniles);  yellowish,  crescent-shaped, 
pigment  patch  in  posterior  part  of  iris,  sometimes  obscure 
due  to  condition  of  preservation.  Aphakic  space  well  devel- 
oped with  its  horizontal  space  about  1.5  times  as  large  as 
width  of  anterior  part  of  iris. 

Upper  end  of  pectoral  fin  base  well  below  midpoint  be- 
tween lateral  line  and  ventral  margin  of  body;  its  lower  end 
above  ventral  margin  of  body  by  a distance  more  than  length 
of  its  base.  Dorsal  fin  origin  behind  center  of  body.  Ventral 
fin  base  below  or  slightly  in  advance  of  posterior  end  of  dorsal 
fin  base.  Adipose  fin  base  above  center  of  anal  fin  base  or 
slightly  in  front  of  it. 

Skin  around  adipose  fin  base  and  caudal  fin  base  pig- 
mented. Ventral  fin  base  slightly  pigmented.  Body  color  in 
preserved  specimen  brown. 

SIZE.  The  largest  specimen  examined  was  266  mm  SL, 
with  mature  ovaries  in  September. 


DISTRIBUTION.  Nansenia  crassa  is  found  in  the  Cali- 
fornia Current  region  between  25°N  and  35°N,  where  its 
distribution  overlaps  that  of  N.  Candida  off  mid-  and  south- 
ern California.  It  also  occurs  in  the  eastern  tropical  Pacific 
along  the  equator  between  8°N  and  7°S  west  to  at  least  1 26°W 
(Ahlstrom,  1971,  1972).  The  pattern  of  occurrence  of  adult 
specimens  is  associated  with  slope  waters  off  California,  sug- 
gesting that  the  adults  there  are  benthopelagic  life  on  the 
continental  or  insular  slope.  Larvae  and  juveniles  are  pelagic 
in  the  upper  200-m  layer  and  are  more  broadly  distributed 
than  are  adults.  The  larvae  of  N.  crassa  can  be  distinguished 
from  those  of  N.  Candida.  The  geographical  distributions  of 
the  larvae  of  the  two  species  correspond  largely  to  the  dis- 
tributions of  the  adults,  although  there  is  a larger  area  of 
overlap  in  the  larvae.  Specimens  between  60  mm  and  160 
mm  SL  are  completely  lacking  in  our  material.  The  size  at 
which  the  specimens  begin  to  disappear  from  our  material 
corresponds  to  an  abrupt  change  in  allometric  growth  (Fig. 
20).  At  this  size,  50-60  mm  SL,  N.  crassa  may  change  its 
vertical  distributional  pattern  from  pelagic  to  benthopelagic 
life  and  become  unavailable  to  either  pelagic  micronekton 
nets  or  bottom  trawls  with  coarse  mesh. 

DISCUSSION 

DISTRIBUTION  PATTERN  AND 
ZOOGEOGRAPHY 

There  are  two  types  of  distribution  patterns,  coastal  and 
oceanic,  among  the  species  of  Nansenia. 

Nansenia  ardesiaca  is  restricted  to  slope  waters  associated 
with  insular  or  continental  slopes,  indicating  a pseudo-ocean- 
ic distribution  (Krefft,  1974).  Sampling  data  suggest  that  this 
species  is  pelagic  during  larval  and  juvenile  stages,  but  be- 
comes benthopelagic  as  an  adult.  A distribution  pattern  sim- 
ilar to  that  of  N.  ardesiaca  is  not  uncommon  among  slope 
water  species  of  myctophids  such  as  Diaphus  watasei,  D. 
garmani,  D.  suborbitalis,  Benthosema  fibulatum,  and  B. 
pterotum  (Nafpaktitis,  1978;  Gjosaeter,  1981).  These  myc- 
tophids are  distributed  in  Southeast  Asian  Seas  and  extend 
to  off  southern  Japan  and  westward  in  the  Indian  Ocean  to 
the  Mozambique  Channel,  although  few  samples  exist  from 
the  slope  waters  off  India.  Of  the  remaining  1 1 species,  three 
(N.  crassa,  N.  oblita,  and  N.  Candida)  show  an  intermediate 
pattern  of  distribution  between  coastal  and  oceanic  (Figs.  2 1 , 
22).  Off  southern  California  N.  crassa  has  a coastal  distri- 
bution, however  the  larvae  are  oceanic  in  the  eastern  tropical 
Pacific  (Ahlstrom,  1971,  1972).  The  other  nine  species  are 
all  oceanic. 

The  1 3 species  may  also  be  grouped  into  cold-water  forms, 
four  species;  warm-water  forms,  seven  species  (sensu  John- 
son, 1982);  and  transitional  forms,  two  species. 

Of  the  four  cold-water  species,  N.  Candida  occurs  only  in 
the  subarctic  eastern  Pacific.  Kanayama  and  Amaoka  (1983) 
described  Nansenia  sanrikuensis,  the  validity  of  which  re- 
quires further  confirmation,  from  the  subarctic  waters  of  the 
western  Pacific  off  Japan.  Species  assemblages  have  been 
recognized  in  the  myctophids  of  both  the  eastern  and  western 


", 


Contributions  in  Science,  Number  352 


Kawaguchi  and  Butler:  Fishes  of  the  Genus  Nansenia  19 


Figure  22.  Distributions  of  specimens  examined  of  six  species  of  Nansenia.  A single  symbol  may  indicate  more  than  one  record. 


parts  of  the  subarctic  Pacific,  which  includes  Diaphus  theta, 
Lampanyctus  ritteri,  L.  jordani,  Stenobrachius  leucopsarus, 
S.  nannochir,  Symbo/ophorus  calif orniensis,  and  Tarleton- 
beania  crenularis  (Wisner,  1974).  Clinal  variation  occurs  in 
the  eye  diameter  of  D.  theta  (Nafpaktitis,  1978;  Kawaguchi 
and  Shimizu,  1978)  and  in  the  shape  of  the  caudal  luminous 
scales  of  S.  californiensis  (Wisner,  1974  and  pers.  commun.) 
A similar  species-subspecies  problem  exists  in  Tarleton- 
beania  (Wisner,  1959;  Becker,  1966).  Definitive  comparative 
examination  of  these  species  assemblages  based  on  materials 
not  only  from  both  sides  of  the  subarctic  Pacific,  but  from 
the  central  area  is  needed.  At  present,  it  is  not  possible  to 
establish  either  a clear  break  in  the  distribution  of  the  eastern 
and  western  populations  (or  species)  or  clinal  variations  in 
characters. 

Other  cold-water  species,  N.  tenera  and  N.  groenlandica, 
belong  to  a subpolar-temperate  group  based  on  the  system 
of  distribution  patterns  recognized  for  Atlantic  mid  water  fishes 
by  Backus  et  al.  (1977).  These  two  species  occur  in  both 
hemispheres,  although  N.  groenlandica  in  the  southern  hemi- 
sphere is  restricted  to  the  subpolar-temperate  region  of  the 
eastern  South  Pacific.  However,  some  bipolar  species  of  the 
subpolar-temperate  group  in  the  Atlantic  such  as  Lampa- 
nyctus macdonaldi  (Wisner,  1974;  Backus  et  al.,  1977)  and 
Gonostoma  bathyphilum  (Mukhacheva,  1972)  are  found 
westward  to  the  eastern  South  Pacific  and  occur  north  to 
Chile.  Therefore  additional  sampling  might  establish  the 
presence  of  N.  groenlandica  in  the  subantarctic  Atlantic.  Of 
the  remaining  cold-water  species,  N.  antarctica  is  probably 


circumpolar  along  the  Antarctic  convergence,  although  col- 
lections are  needed  in  the  Indian  Ocean  sector. 

Two  of  the  seven  warm -water  species,  N.  atlantica  and  N. 
megalopa,  occur  only  in  the  eastern  tropical  Atlantic.  Ac- 
cording to  the  Atlantic  faunal  regions  recognized  by  Backus 
et  al.  (1977),  both  species  inhabit  the  Guinean  Province  of 
Atlantic  Tropical  Region  (Fig.  21).  The  distribution  of  N. 
atlantica  appears  to  be  restricted  to  the  more  productive  area 
near  the  Mauritanian  upwelling  and  to  just  off  the  Guinean 
coast.  The  other  two  warm-water  species,  N.  longicauda  and 
N.  pelagica,  are  widely  distributed  in  the  Atlantic  and  Pacific 
oceans  with  the  former  restricted  to  the  subtropical  region 
between  20°N  and  40°N  and  the  latter  to  the  tropical  region 
between  20°N  and  20°S  in  the  Atlantic  and  between  30°N 
and  20°S  in  the  Pacific.  Nansenia  ahlstromi  is  restricted  to 
the  central  Eastern  North  Pacific.  There  is  nothing  to  add  to 
the  previous  discussion  of  the  distribution  of  the  remaining 
two  warm-water  species,  N.  oblita  and  N.  ardesiaca. 

BODY  COLOR 

There  have  been  many  descriptions  of  the  body  color  of 
Nansenia,  which  suggest  two  patterns,  silvery  and  colored. 
During  the  course  of  this  study,  however,  it  has  been  observed 
that  the  silvery  color  pattern  is  variable,  depending  upon 
preservative;  it  is  easily  lost  in  formalin.  For  example,  the 
type  of  N.  ardesiaca  (Fig.  1 3a),  which  is  now  dark  colored, 
was  originally  described  as  “color  brilliant  silvery  . . . scales 
of  a bright,  silvery  appearance”  (Jordan  and  Evermann,  1 896). 


20  Contributions  in  Science,  Number  352 


Kawaguchi  and  Butler:  Fishes  of  the  Genus  Nansenia 


Some  specimens  examined  (Fig.  1 3b)  still  bear  complete  gua- 
nine after  more  than  50  years  of  preservation  in  ethanol  in 
a dark  place.  But  these  specimens  are  usually  so  soft  that 
careful  handling  is  necessary,  indicating  that  they  were  not 
fixed  well  in  formalin. 

INTERRELATIONSHIPS 

Various  character  states  studied  here  are  shared  indepen- 
dently among  13  species  and  it  is  difficult  to  discuss  the 
relationships  within  the  species  of  Nansenia.  Although  the 
key  separates  species  on  the  number  of  branchiostegal  rays, 
loss  of  a branchiostegal  ray  could  have  occurred  more  than 
once.  It  is  also  possible  to  separate  the  species  of  this  genus 
into  three  groups  based  on  morphology,  mainly  of  the  caudal 
peduncle.  With  the  exception  of  N.  antarctica,  the  species  of 
Nansenia  are  separable  into  the  following  three  distinctive 
groups  based  on  the  ratio  of  caudal  peduncle  length  to  caudal 
peduncle  depth  (CPL/CPD,  Tables  1 and  2): 

Stubby-tailed  group  CPL/CPD  = 1.0-1. 5 

Intermediate  group  CPL/CPD  = 1.7-2. 3 

Slender-tailed  group  CPL/CPD  = 2.7-4. 1 

The  ratios  are  not  overlapping.  The  stubby-tailed  group  in- 
cludes four  species:  N.  ahlstromi  (Br  3),  N.  oblita  (Br  4),  N. 
atlantica  (Br  4),  and  N.  pe/agica  (Br  4).  The  slender-tailed 
group  includes:  N.  groenlandica  (Br  3),  TV.  tenuicauda  (Br  4), 
N.  megalopa  (Br  4),  and  N.  longicauda  (Br  4).  Of  the  re- 
maining five,  four  species,  N.  Candida  (Br  3),  N.  crassa  (Br 
4),  N.  ardesiaca  (Br  4),  and  N.  tenera  (Br  4),  are  included  in 
the  intermediate  group.  Nansenia  antarctica  (Br  3)  shows 
remarkable  allometric  growth  in  the  caudal  peduncle;  the 
range  of  the  CPL/CPD  ratio  for  specimens  less  than  1 10  mm 
SL  was  2. 5-3.0;  but  the  ratio  for  specimens  larger  than  1 10 
mm  SL  was  2.0-2. 5.  This  overlaps  the  range  between  inter- 
mediate and  slender-tailed  groups. 

No  correspondence  is  recognized  between  groupings  based 
on  the  number  of  branchiostegal  rays  or  morphology.  This 
suggests  that  natural  groups  cannot  be  defined  by  branchio- 
stegal ray  counts  alone. 

ACKNOWLEDGMENTS 

We  thank  the  late  E.H.  Ahlstrom  whose  interest  in  and  en- 
thusiasm for  fish  systematics  were  largely  responsible  for  this 
study.  We  are  greatly  indebted  to  H.G.  Moser  and  his  col- 
leagues of  the  Southwest  Fisheries  Center  for  their  continuous 
help  and  kindness  during  the  course  of  this  study.  The  fol- 
lowing people  and  their  institutions  allowed  us  to  examine 
the  materials  in  their  care:  E.  Bertelsen  (ZMUC);  G.  Krefft 
and  A.  Post  (ISH);  R.J.  Lavenberg  (LACM);  R.H.  Rosenblatt 
(SIO);  W.G.  Pearcy  (OSU);  S.H.  Weitzman  (USNM);  R.K. 
Johnson  (FMNH);  W.N.  Eschmeyer  (CAS);  G.H.  Burgess 
(UF);  O.  Okamura  (KU).  We  would  also  like  to  thank  D.M. 
Cohen  (LACM),  R.H.  Rosenblatt  (SIO),  and  D.L.  Stein  (OSU) 
for  reading  the  manuscript  and  for  their  many  helpful  com- 
ments. The  authors  thank  I.  Barrett,  director  of  the  Southwest 
Fisheries  Center  and  Professor  Emeritus  R.  Marumo  of  Uni- 
versity of  Tokyo  for  giving  them  a chance  to  complete  this 
study. 


Partial  financial  support  from  the  Japanese  Ministry  of 
Education,  special  project  research  “Studies  on  biological 
processes  in  the  sea  and  technical  development  of  marine 
resources”  (No.  57108013)  is  also  acknowledged. 


LITERATURE  CITED 

Abe,T.  1976.  Notes  on  some  fishes  collected  by  the  fisheries 
research  vessel  “Kaiyo-maru”  in  the  South  China  Sea. 
IV.  Bulletin  of  the  Biogeographical  Society  of  Japan  31(4): 
27-31. 

Ahlstrom,  E.H.  1971.  Kinds  and  abundance  of  fish  larvae 
in  the  eastern  tropical  Pacific,  based  on  collections  made 
on  EASTROPAC  I.  Fishery  Bulletin  (U.S.)  69:3-77. 

. 1972.  Kinds  and  abundance  of  fish  larvae  in  the 

eastern  tropical  Pacific  on  the  second  multivessel  EAS- 
TROPAC survey,  and  observations  on  the  annual  cycle 
of  larval  abundance.  Fishery  Bulletin  (U.S.)  70:1  153- 
1242. 

Ahlstrom,  E.H.,  H.G.  Moser,  and  D.M.  Cohen.  1984.  On- 
togeny and  systematics  of  fishes  Suborder  Argeninoidei. 
In  Moser,  H.G.,  W.J.  Richards,  et  al..  Ontogeny  and 
systematics  of  fishes.  Spec.  Publ.  No.  1,  Amer.  Soc.  Ich- 
thyol.  Herpetol.  (in  press). 

Backus,  R.H.,  J.E.  Craddock,  R.L.  Haedrich,  and  R.H.  Rob- 
ison. 1977.  Atlantic  mesopelagic  zoogeography.  Mem- 
oir Sears  Foundation  for  Marine  Research,  I (part  7): 
266-287. 

Becker,  V.E.  1966.  Slender  tailed  myctophids  (genera  Lo- 
weina,  Tarletonbeania,  Gonichthys,  and  Centrobran- 
chus)  of  the  Pacific  and  Indian  oceans,  systematics  and 
distribution.  Pages  10-78  in  Rass,  T.S.,  ed.  (L.  Penney, 
E.  Roden,  and  E.  Roifer,  trans.),  Fishes  of  the  Pacific 
and  Indian  oceans,  biology  and  distribution.  Israel  Pro- 
gram for  Scientific  Translation,  Jerusalem. 

Belloti,  C.  1888.  Note  ittiologiche.  Osservazione  fatte  sulla 
collezione  ittiologica.  Atti  della  Societe  Italiana  Scienza 
Naturali  31:213-229. 

Blache,  J.,  and  M.  Rossignol.  1962.  III.  Nansenia  atlantica 
nov.  sp.  (Teleostei,  Clupeiformes,  Opisthoproctoidei, 
Microstomidae),  Poisson  bathypelagique  nouveau  du 
Golfe  de  Guinee.  Cahiers  O.R.S.T.O.M.  Serie  Oceano- 
graphie  (2):  103-106. 

Chapman,  W.M.  1942.  The  osteology  and  relationships  of 
the  Argentinidae,  a family  of  oceanic  fishes.  Journal  of 
the  Washington  Academy  of  Sciences  32:104-1 17. 

. 1943.  The  osteology  and  relationships  of  the  bathy- 

pelagic  fishes  of  the  genus  Bathylagus  Gunther,  with 
notes  on  the  systematic  position  of  Leuroglossus  stilbius 
Gilbert  and  Therobromus  callorhini  Lucas.  Journal  of 
the  Washington  Academy  of  Sciences  33(5):  147-160. 

. 1948.  The  osteology  and  relationships  of  the  Mi- 
crostomidae, a family  of  oceanic  fishes.  Proceedings  of 
the  California  Academy  of  Sciences,  Ser.  4,  26(1):  1-22. 

Cohen,  D.M.  1958.  Nansenia  Candida,  a new  species  of 
argentinid  fish  from  the  north  Pacific,  with  notes  on  the 
other  species  of  Nansenia.  Stanford  Ichthyological  Bul- 
letin 7:52-57. 


Contributions  in  Science,  Number  352 


Kawaguchi  and  Butler:  Fishes  of  the  Genus  Nansenia  21 


. 1964.  Suborder  Argentinoidea.  Memoir  Sears 

Foundation  for  Marine  Research,  New  Flaven  1(4):  1- 
70. 

Facciola,  L.  1887.  Sull  esistenza  di  due  forme  diverse  di 
Microstoma  nel  mar  di  Messina.  Naturalista  Siciliano, 
Palermo  6:193-197. 

Follet,  M.I.,  and  D.M.  Cohen.  1958.  Request  for  a ruling 
as  to  the  species  to  be  accepted  as  the  type  species  of  the 
nominal  genus  “ Bathylagus”  Gunther  (A.C.L.G.),  1878 
(class  Pisces).  Bulletin  of  Zoological  Nomenclature  16(2): 
73-78. 

Fowler,  H.W.  1934.  Descriptions  of  new  fishes  obtained 
1907  to  1910  chiefly  in  the  Philippine  Islands  and  ad- 
jacent seas.  Proceedings  of  the  Academy  of  Natural  Sci- 
ences, Philadelphia  85:233-367. 

Gilchrist,  J.D.J.  1922.  Deep-sea  fishes  procured  by  the  SS 
‘Pickle’  (Part  I).  Report  of  the  Fisheries  Marine  Biolog- 
ical Survey,  Republic  of  South  Africa,  2 (Special  Report 
3):4 1 —79,  pis.  VII-XIII. 

Gjosaeter,  J.  1981.  Growth,  production  and  reproduction 
of  the  myctophid  fish,  Benthosema  glaciate,  from  west- 
ern Norway  and  adjacent  seas.  Fiskerdirektorates  Skrif- 
ter.  Serie  Havundersokelser  17:79-108. 

Hubbs,  C.L.,  and  K.F.  Lagler.  1964.  The  fishes  of  the  Great 
Lakes  region.  University  of  Michigan  Press,  Ann  Arbor. 
213  pp. 

Johnson,  R.K.  1982.  Fishes  of  the  families  Euermannel- 
lidae  and  Scopelarchidae:  Systematics,  morphology,  in- 
terrelationships, and  zoogeography.  Fieldiana  12:1-252. 

Jordan,  D.S.,  and  B.W.  Evermann.  1896.  The  fishes  of 
North  and  Middle  America,  descriptive  catalogue  of  the 
species  of  fish-like  vertebrates  found  in  the  waters  of 
North  America,  north  of  the  isthmus  of  Panama.  United 
States  National  Museum,  Bulletin  (47),  Part  1,  lx  + 1 240 
pp. 

Jordan,  D.S.,  and  W.F.  Thompson.  1914.  Record  of  the 
fishes  obtained  in  Japan  in  1911.  Memoirs  of  the  Car- 
negie Museum  6:205-313. 

Kanayama,  T.,  and  K.  Amaoka.  1983.  Nansenia  sanri- 
kuensis  (Kurosake-iwashi).  Pages  77-79  in  Amaoka  et 
al.  (eds.).  Fishes  from  the  northeastern  sea  of  Japan. 
Japan  Fisheries  Resource  Conservation  Association. 

Kawaguchi,  K.,  and  H.  Shimizu.  1978.  Taxonomy  and 
distribution  of  the  lantemfishes,  genus  Diaphus  (Pisces, 
Myctophidae)  in  the  western  Pacific,  eastern  Indian 
Oceans  and  the  Southeast  Asian  Seas.  Bulletin  of  the 
Ocean  Research  Institute,  University  of  Tokyo,  No.  10, 
145  pp. 


Krefft,  G.  1974.  Investigations  on  midwater  fish  in  the 
Atlantic  Ocean.  Berichte  der  Deutschen  Wissenshaft- 
lichen  Kommission  fur  Meeresforschung  23:226-254. 

Kuroda,  N.  1947.  Additional  list  of  fishes  in  Suruga  Bay 
off"  Sizu-ura  (No.  8)  Seibutsu  (Life)  (1):25— 3 1 . 

. 1951.  A nominal  list  with  distribution  of  the  fishes 

of  Suraga  Bay,  inclusive  of  the  fresh  water  species  found 
near  the  coast.  Japanese  Journal  of  Ichthyology  1(5): 
314-338,  376-394. 

Lavenberg,  R.K.  1965.  A new  species  of  microstomatid 
fish  ( Nansenia ) from  California  waters.  Copeia  1965(3): 
281-385. 

Matsubara,  K.  1955.  Fish  morphology  and  hierarchy,  Parts 
I— III.  Ishi-zaki  Shoten,  Tokyo.  1605  pp. 

Mukhacheva,  V.A.  1972.  On  systematics,  distribution  and 
biology  of  the  Gonostoma  species  (Pisces,  Gonostomat- 
idae).  Trudy  Instituta  Okeanologii  Akademiya  Nauk, 
SSSR  93:205-249. 

Nafpaktitis,  B.G.  1978.  Systematics  and  distribution  of  lan- 
temfishes of  the  genera  Lobianchia  and  Diaphus  (Myc- 
tophidae) in  the  Indian  Ocean.  Natural  History  Museum 
of  Los  Angeles  County,  Bulletin  No.  30,  92  pp. 

Parin,  N.V.,  and  G.A.  Golovan.  1976.  Pelagic  deep-sea 
fishes  of  the  families  characteristic  of  the  open  ocean 
collected  over  the  continental  slope  off  west  Africa.  Tru- 
dy Instituta  Okeanologii,  Akademiya  Nauk,  SSSR  104: 
250-276. 

Reinhardt,  J.C.H.  1840.  [Gronlands  Fiske]:8-10.  In  Or- 
sted,  H.C.,  Oversikt  Konelige  Danske  Videnskabemes 
Selskabs  Forhandlinger  (1839):  1-44. 

Sarato,  C.  1890.  Notes  sur  les  poissons  de  Nice.  (Reprint: 
1-3.)  Nice. 

Schmidt,  E.J.  1918.  Argentinidae,  Microstomidae,  Opis- 
thoproctidae,  Mediterranean  Odontostomidae.  Report 
on  the  Danish  Oceanographical  Expeditions  1908-10  to 
the  Mediterranean  and  adjacent  seas,  2.  Biology  (A5): 
1-40. 

Tanaka,  S.  1911.  Figures  and  descriptions  of  the  fishes  of 
Japan.  Vol.  1.  Dai-ichi  shoin,  Tokyo.  18  pp. 

Wisner,  R.L.  1959.  Distribution  and  differentiation  of  the 
North  Pacific  myctophid  fish,  Tarletonbeania  taylori. 
Copeia  1959(1):  1-7. 

. 1974.  The  taxonomy  and  distribution  of  lantem- 
fishes (Family  Myctophidae)  of  the  eastern  Pacific  Ocean. 
NORDA  Report  3,  Department  of  the  Navy  (U.S.).  229 

pp. 

Accepted  28  March  1984. 


22  Contributions  in  Science,  Number  352 


Kawaguchi  and  Butler:  Fishes  of  the  Genus  Nansenia 


INSTIL! CHONS  FO ft  AtprpMS 


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KENTRIOBON  O&SCUkUS  (KEI.I,OG€?  1931),  A FOSSIL  DOLPHIN 
(MAMMALIA:  KENll?i.ODONlIiT]f4!E;):  FROM  THE  MIOCENE 
SHARKTOOTH  HILL  BONEBED  IN  CAIJFORNIA 


Lawrence  G.  Barnes  and  Edward  MSfclseil 


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sit  Contributions  in  'Science,  a misce 


inai  research  in  the  life  and.  earth  sciences. 

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SCIENTIFIC  PUBLICATIONS  COMMIT! EE 


Craig  C.  Black,  Museum  Director 
Donald  Chaput 
Dame!  M Cohen,  Commit 
John  M.  Hanr;s 
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KENTRIODON  OBSCURUS  (KELLOGG,  1931),  A FOSSIL  DOLPHIN 
(MAMMALIA:  KENTRIODONTIDAE)  FROM  THE  MIOCENE 
SHARKTOOTH  HILL  BONEBED  IN  CALIFORNIA 

Lawrence  G.  Barnes'  and  Edward  Mitchell1 2 


ABSTRACT.  The  fossil  odontocete  genus  Kentriodon  Kellogg,  1927, 
is  the  type  genus  of  the  delphinoid  family  Kentriodontidae,  a family 
which  has  no  living  representatives.  The  genus  Kentriodon  has  pre- 
viously included  only  one  named  species,  a small  dolphin  called 
Kentriodon  pernix  Kellogg,  1927,  from  the  Middle  Miocene  Calvert 
Formation  in  Maryland  and  Virginia  on  the  east  coast  of  North 
America.  Some  bones  from  Japan  and  California  have  previously 
been  tentatively  referred  to  the  genus.  We  have  concluded  that  the 
species  Grypolithax  obscura  Kellogg,  1931,  heretofore  known  only 
by  isolated  periotics  from  the  Middle  Miocene  Sharktooth  Hill  Bone- 
bed  in  California,  belongs  in  Kentriodon.  The  genus  Grypolithax 
Kellogg,  1931,  is  therefore  a junior  synonym  of  Kentriodon  Kellogg, 
1927.  A second  species  named  on  isolated  periotics  from  the  Shark- 
tooth Hill  Bonebed,  Grypolithax pavida  Kellogg,  1 93 1 , is  a synonym 
of  G.  obscura.  The  appropriate  binomen  for  the  species  is  Kentriodon 
obscurus  (Kellogg,  1931).  We  assign  to  this  species  a partial  skull 
and  additional  isolated  periotics  from  the  same  bonebed. 

Other  isolated  periotics  from  rocks  stratigraphically  below  the 
Sharktooth  Hill  Bonebed  in  California  resemble  those  of  Kentriodon 
spp.  and  a closely  related  kentriodontid,  Delphinodon  dividum  True, 
1912,  which  was  also  originally  described  from  the  Calvert  For- 
mation. Some  of  these  have  been  referred  to  in  previous  literature, 
but  are  illustrated  and  described  here  for  the  first  time. 

These  newly  reported  specimens  from  California  reinforce  some 
previous  correlations  between  the  upper  part  of  the  Round  Mountain 
Silt  in  California  and  the  Calvert  Formation  in  Maryland  and  Vir- 
ginia. They  also  provide  additional  indications  of  a general  pattern 
of  generic  cosmopolitanism  and  specific  endemism  among  the  Mio- 
cene odontocetes  of  the  North  Atlantic  and  North  Pacific  Oceans. 

INTRODUCTION 

Fossil  odontocetes  of  the  genus  Kentriodon  Kellogg,  1927, 
are  small  Miocene  dolphins  in  the  extinct  family  Kentri- 
odontidae. Kellogg  (1928:33,  68)  assigned  the  genus  to  the 
family  Delphinidae.  Slijper  later  (1936:556)  named  a new 
subfamily  Kentriodontinae,  which  he  placed  within  the  Del- 
phinidae, to  include  Kentriodon  as  well  as  Delphinodon  div- 
idum True,  1912,  another  Miocene  dolphin.  Barnes  (1978) 
altered  the  context  and  rank  of  Slijper’s  family  group  name 
by  recognizing  the  family  Kentriodontidae  within  the  super- 
family Delphinoidea.  He  recognized  three  subfamilies  (Kam- 
pholophinae,  Kentriodontinae,  and  Lophocetinae)  within  the 
Kentriodontidae,  and  the  species  included  in  this  family  are 
now  known  by  described  fossils  from  Europe,  Japan,  New 
Zealand,  and  both  the  east  and  west  coasts  of  North  America. 
The  importance  of  this  family  and  its  relationships  to  some 
other  odontocete  families  was  commented  on  by  Barnes. 
He  stated  that  kentriodontids  might  comprise  the  group  from 
which  other  living  families  of  delphinoids,  including  modem 
dolphins  in  the  Delphinidae,  have  evolved,  and  they  might 
also  be  expected  to  occur  widely  in  rocks  of  appropriate  age 

Contributions  in  Science,  Number  353,  pp.  1-23 
Natural  History  Museum  of  Los  Angeles  County,  1984 


around  the  world.  Our  interests  in  delphinoid  evolution  and 
in  the  Middle  Miocene  cetaceans  in  the  Sharktooth  Hill  Local 
Fauna  in  California  have  led  us  to  the  present  study. 

The  type  species  of  Kentriodon,  K.  pernix  Kellogg,  1927, 
was  named  on  the  basis  of  two  skulls,  one  of  which  was 
collected  with  earbones  and  an  articulated  partial  postcranial 
skeleton  (the  holotype),  of  Middle  Miocene  age  from  the 
Calvert  Formation  in  Maryland.  Until  the  present  study,  this 
genus  has  had  assigned  to  it  only  this  one  named  species, 
however,  some  references  in  the  paleontologic  literature  have 
suggested  the  former  presence  of  the  genus  in  the  North  Pa- 
cific Ocean.  The  genus  Kentriodon  was  questionably  iden- 
tified from  Miocene  rocks  in  Japan  by  Shikama,  Hasegawa, 
and  Otsuka  (1973;  also  cited  in  Okazaki,  1976:25),  but  spec- 
imens documenting  this  identification  have  not  been  de- 
scribed. Bames  (1976:326)  identified  as  cf.  Kentriodon  and 
as  a related  kentriodontid,  cf.  Delphinodon  dividum  True, 
1912,  some  isolated  periotics  of  early  Middle  Miocene  age 
from  the  lower  part  of  the  Round  Mountain  Silt  in  Kern 
County,  California.  The  source  of  these  is  stratigraphically 
lower  than  the  Middle  Miocene  Sharktooth  Hill  Bonebed. 
Other  kentriodontid  genera  have  been  recognized  based  on 
skulls  from  Miocene  rocks  in  Europe  and  California  (Kellogg, 
1925:4—6;  Bames,  1976,  1978),  and  from  late  Oligocene  rocks 
in  New  Zealand  (Fordyce,  1980:328). 

Kellogg  (1931)  prepared  a preliminary  description  of  the 
mammals  from  the  Sharktooth  Hill  Bonebed  that  comprise 
a part  of  what  we  now  call  the  Sharktooth  Hill  Local  Fauna 
(Mitchell,  1965:33;  Mitchell  and  Tedford,  1973:  fig.  3;  Bames, 
1 976:326-327).  In  this  study,  Kellogg  named  ten  new  species 
of  small  odontocetes  based  solely  on  isolated  periotics  from 
the  bonebed.  For  many  years  the  true  identities  and  rela- 
tionships of  these  species  have  remained  unknown.  The 
problem  of  disparate  skeletal  parts  has  in  some  instances 
precluded  objective  morphologic  and  taxonomic  compari- 
sons between  the  odontocetes  from  the  Sharktooth  Hill  Bone- 
bed and  other  taxa  (even  some  studied  by  Kellogg  himself) 
known  by  skulls  and  skeletons  from  elsewhere  in  the  world. 
For  example,  Bames  (1978)  showed  that  Liolithax  kernensis 
Kellogg,  1931,  from  the  Sharktooth  Hill  Bonebed  is  conge- 
neric with  Lophocetus  pappus  Kellogg,  1955,  from  the  Cal- 
vert Formation  in  Maryland. 

In  the  present  study,  we  show  that  Grypolithax  obscura 
Kellogg,  1931,  from  the  Sharktooth  Hill  Bonebed  is  conge  - 


1.  Section  of  Vertebrate  Paleontology,  Natural  History  Museum 
of  Los  Angeles  County,  Los  Angeles,  California  90007. 

2.  Arctic  Biological  Station,  Fisheries  and  Oceans,  Ste.  Anne  de 
Bellevue,  Quebec,  Canada  H9X  3R4;  and  Research  Associate,  Nat- 
ural History  Museum  of  Los  Angeles  County. 


ISSN  0459-8113 


neric  with  Kentriodon  pernix.  Another  species  Kellogg  named 
from  the  bonebed,  G.  pavida  Kellogg,  1931,  is  synonymous 
with  G.  obscura  and  the  properly  emended  binomen  for  the 
species  from  California  is  Kentriodon  obscurus  (Kellogg,  1931). 
We  refer  a partial  skull  and  several  more  isolated  periotics 
from  the  Sharktooth  Hill  Bonebed  to  K.  obscurus,  and  with 
these  we  have  been  able  to  more  fully  characterize  the  species 
and  compare  it  with  K.  pernix.  We  also  describe  and  illustrate 
the  specimens  of  kentriodontids  that  Barnes  (1976)  men- 
tioned from  stratigraphically  lower  levels  of  the  Round 
Mountain  Silt  in  California. 

MATERIALS  AND  METHODS 
INSTITUTIONAL  ACRONYMS 

The  specimens  examined  in  the  course  of  this  study  are  in 
the  collections  of  the  following  scientific  institutions:  Cali- 
fornia Academy  of  Sciences,  San  Francisco,  California  (CAS); 
Natural  History  Museum  of  Los  Angeles  County,  Los  An- 
geles, California  (LACM);  University  of  California  Museum 
of  Paleontology,  Berkeley,  California  (UCMP);  National  Mu- 
seum of  Natural  History,  Smithsonian  Institution,  Washing- 
ton, D.C.  (USNM). 

Institutional  locality  numbers  are  given  for  specimens  where 
appropriate.  Qualified  investigators  may  obtain  precise  lo- 
cality information  by  contacting  the  appropriate  institutions. 

COMPARATIVE  MATERIAL 

We  have  made  extensive  reference  to  and  comparisons  with 
the  type  species  of  Kentriodon,  K.  pernix  from  the  Middle 
Miocene  Calvert  Formation  in  Maryland.  We  believe  that 
the  holotype  (USNM  8060)  of  K.  pernix,  which  was  collected 
from  Zone  5 relatively  low  in  the  Calvert  Formation,  and 
Kellogg’s  original  (1927a)  published  referred  skull  (USNM 
1 0670)  from  Zone  3 slightly  lower  in  the  formation,  represent 
the  same  species.  These  are  the  only  skulls  of  K.  pernix  that 
have  been  described  in  the  literature. 

We  have  identified  one  other  small  skull  (USNM  21027) 
as  K.  pernix.  It  was  found  at  Plum  Point,  Maryland,  where 
Zone  6 of  the  Calvert  Formation  (also  relatively  low  in  the 
rock  unit)  is  exposed  at  sea  level  (see  Clark,  Shattuck,  and 
Dali,  1 904:  pi.  5).  These  three  specimens,  therefore,  comprise 
our  current  concept  of  the  species,  but  provide  only  limited 
information  on  individual  and  ontogenetic  variation  within 
the  species. 

There  are  several  additional  fossils  of  Kentriodon  from  the 
Calvert  Formation  in  Virginia  and  Maryland  in  the  USNM 
collections  that  are  undescribed  in  the  scientific  literature. 
Some  of  these  are  significantly  different  from  both  the  ho- 
lotype and  referred  specimens  of  K.  pernix.  Most  are  from 
zones  higher  in  the  Calvert  Formation,  and  are  therefore 
geologically  younger  than  the  holotype  and  referred  speci- 
mens of  K.  pernix.  We  conclude  that  one  or  more  species  in 
addition  to  K.  pernix  are  represented  by  these  additional 
specimens,  but  a detailed  variability  and  taxonomic  study  of 
Kentriodon  from  the  Calvert  Formation  is  beyond  the  scope 
of  the  present  study.  In  the  absence  of  such  a study  of  these 


other  Kentriodon  specimens,  we  made  some  observations  on 
them  that  have  influenced  our  diagnoses  and  comparisons. 
Additionally,  we  have  illustrated  some  of  the  periotics  (Figs. 
8-9)  to  demonstrate  variable  characters  within  the  genus. 

TERMINOLOGY 

The  morphology  of  a kentriodontid  periotic  is  shown  in  Fig. 

1 . The  terminology  used  is  derived  or  adapted  from  Denker 
(1902),  Boenninghaus  (1904),  and  Kellogg  (1928).  Cranial 
terminology  is  derived  from  Kellogg  (1927a)  and  Fraser  and 
Purves  (1960).  Where  we  employ  family  group  names  with 
different  rank  than  originally  proposed,  we  cite  the  author  of 
the  emended  rank  following  the  original  author. 

Statistical  analysis  follows  Simpson,  Roe,  and  Lewontin 
(1960). 

SYSTEMATICS 

Class  Mammalia  Linnaeus,  1758 

Order  Cetacea  Brisson,  1762 

Suborder  Odontoceti  Flower,  1867 

Superfamily  Delphinoidea  (Gray,  1821) 
Rower,  1864 

Family  Kentriodontidae  (Slijper,  1936) 
Barnes,  1978 

Kentriodontinae  Slijper,  1 936:556;  as  a subfamily  of  the  fam- 
ily Delphinidae. 

Kentriodontidae.  Slijper,  1958:  label  in  fig.  36,  emended  rank 
without  explanation  in  text. 

Kentriodontidae.  Barnes,  1978:3;  emended  rank,  as  a family 
of  the  superfamily  Delphinoidea. 

DISCUSSION.  Barnes  (1978)  stated  that  a lack  of  fossae 
in  bones  of  the  basicranium,  reflecting  a presumed  lack  of 
extensive  development  of  air  sinuses  of  the  middle  ear  air 
sinus  system,  was  a characteristic  feature  of  the  family  Ken- 
triodontidae. Aside  from  the  usual,  primitive  odontocete 
combination  of  peribullary  and  pterygoid  air  sinuses,  we  find 
in  Kentriodon  spp.  osteological  evidence  only  for  a middle 
sinus  adjacent  to  the  glenoid  fossa  and  for  a postorbital  lobe 
of  the  pterygoid  sinus.  We  find  no  fossae  that  would  provide 
evidence  for  a posterior  sinus  in  the  exoccipital,  for  any  si- 
nuses in  the  basioccipital  or  above  the  optic  nerve  (as  in 
phocoenids),  for  a large  orbital  lobe,  or  for  an  anterior  sinus 
extending  onto  the  posterior  part  of  the  palate.  These  are  all 
locations  where  various  modem  odontocetes  have  been  shown 
to  have  air  sinuses  in  their  skulls  (Fraser  and  Purves,  1960). 
The  fossa  for  the  lobe  of  the  pterygoid  sinus  within  the  pter- 
ygoid hamulus  is  characteristically  small  in  Kentriodontidae 
compared  with  Delphinidae,  Phocoenidae,  Ziphiidae  or  Phy- 
seteridae.  No  fossil  kentriodontid  skull  has  yet  been  de- 
scribed with  an  ossified  pterygoid  hamulus  flooring  the  ven- 
tral surface  of  this  sinus  as  in  modem  delphinids,  phocoenids, 
and  monodontids,  and  it  cannot  be  determined  from  the 


2 Contributions  in  Science,  Number  353 


Barnes  and  Mitchell:  Kentriodon  from  California 


specimens  presently  available  whether  or  not  these  animals 
had  an  incomplete  or  non-ossified  hamulus  as  in  modem 
platanistoid  dolphins  (see  Fraser  and  Purves,  1960). 

Subfamily  Kentriodontinae  Slijper,  1936 

Kentriodontinae  Slijper,  1936:556;  as  a subfamily  of  the  fam- 
ily Delphinidae. 

Kentriodontinae.  Bames,  1978:24;  emended  context,  as  a 
subfamily  of  the  family  Kentriodontidae. 

Kentriodon  Kellogg,  1927 

Kentriodon  Kellogg,  1927a:4. 

Grypolithax  Kellogg,  1931:393. 

EMENDED  DIAGNOSIS  OF  GENUS.  A genus  of  the 
subfamily  Kentriodontinae  differing  from  Delphinodon  divi- 
dum  by  having  a skull  with  a longer  rostrum,  the  mesorostral 
gutter  not  roofed  over  by  the  premaxillae  at  the  anterior  end, 
a more  concave  lateral  margin  of  the  supraorbital  process, 
palatal  surfaces  of  maxillae  more  transversely  convex,  and  a 
fossa  for  the  postorbital  lobe  of  the  pterygoid  air  sinus  on 
the  ventral  surface  of  the  frontal;  differing  from  Leptodelphis, 
Microphocaena,  Pithanodelphis,  and  Sarmatodelphis  by  hav- 
ing a more  convex  lateral  margin  of  the  supraorbital  process, 
flat  rather  than  bulbous  or  convex  nasal  bones,  and  more 
widely  separated  posterior  ends  of  the  maxillae  at  the  cranial 
vertex  with  a concomitantly  wider  exposure  of  the  frontals, 
differing  from  Leptodelphis  and  Pithanodelphis  by  having  a 
less  elevated  cranial  vertex;  and  differing  from  Liolithax, 
Delphinodon,  Lophocetus,  and  perhaps  other  described  gen- 
era of  Kentriodontidae  by  having  the  anterior-most  pre- 
maxillary tooth  on  each  side  elongated  into  a small  tusk  and 
pointing  anteriorly  from  the  tip  of  the  rostrum. 

TYPE  SPECIES.  Kentriodon  pernix  Kellogg,  1927;  type 
by  original  monotypy. 

INCLUDED  SPECIES.  Kentriodon  pernix  Kellogg,  1 927; 
and  Kentriodon  obscurus  (Kellogg,  1931),  new  combination. 

Kentriodon  obscurus  (Kellogg,  1931), 
new  combination 

Figures  2-7,  8d-t,  9d-t,  13b,  14b 

Grypolithax  obscura  Kellogg,  1931:394. 

Grypolithax  pavida  Kellogg,  1931:396. 

EMENDED  DIAGNOSIS  OF  SPECIES.  A species  of 
Kentriodon  characterized  by  and  differing  from  K.  pernix  by 
having  skull  with  vertically  short  postorbital  process  of  fron- 
tal, posterior  part  of  alveolar  row  curved  medially  on  palatal 
surface  instead  of  extending  along  lateral  margin  of  rostrum, 
posterior  maxillary  alveoli  directed  more  laterally  than  ven- 
trally,  lateral  margin  of  maxilla  adjacent  to  posterior  part  of 
alveolar  row  thickened  and  squared  off  instead  of  being  thin 
and  rounded,  alveoli  for  maxillary  teeth  averaging  1.5  mm 
in  diameter  instead  of  3 mm,  antorbital  notch  narrow  and 
directed  anteriorly  instead  of  wide  and  directed  anterolat- 
erally,  supraorbital  process  of  frontal  thicker  and  more  arched, 


2 16  13 


6 7 


Figure  1.  Right  periotic  of  a kentriodontid,  cf.  Kentriodon  sp., 
UCMP  83605,  with  anatomical  structures  labeled;  a,  cerebral  or 
dorsal  view;  b,  tympanic  or  ventral  view,  natural  size.  1— anterior 
process,  2 — aquaeductus  cochleae,  1— aquaeductus  Fallopii.  4 — 
aquaeductus  vestibuli,  5— canal  for  facial  nerve,  6— cochlear  portion, 
1 —fenestra  ovalis,  8— fenestra  rotunda,  9— foramen  cent  rale,  10  — 
foramen  singulare,  11— fossa  for  head  of  the  malleus,  12  —fossa 
incudis,  1 3 — internal  acoustic  meatus,  14  — posterior  articular  facet 
for  tympanic  bulla,  1 5 — posterior  process,  16  — tractus  spiralis  fora- 
minosus. 

medial  margins  of  dorsal  premaxillary  surfaces  on  proximal 
part  of  rostrum  between  antorbital  notches  flat  instead  of 
inclined  medially  toward  mesorostral  gutter,  palatine  bone 
and  pterygoid  sinus  fossa  extended  farther  anteriorly  onto 
palate  instead  of  ending  at  level  of  antorbital  processes,  post- 
orbital lobe  of  pterygoid  sinus  of  middle  ear  air  sinus  system 
occupying  large  fossa  on  ventral  surface  of  supraorbital  pro- 
cess of  frontal,  internal  acoustic  meatus  of  periotic  more 
elliptical  and  less  circular  in  shape. 

HOLOTYPE.  CAS  4349,  right  periotic,  collected  by 
Charles  Morrice  in  1924. 

TYPE  LOCALITY.  CAS  locality  905,  Sharktooth  Hill 
Bonebed,  Kern  County,  California. 

PARATYPE.  CAS  4347,  left  periotic  collected  by  Charles 
Morrice  in  1924  from  CAS  locality  905. 

REFERRED  SPECIMENS  FROM  THE  SHARK- 
TOOTH  HILL  BONEBED.  LACM  21256,  an  incomplete 


Contributions  in  Science,  Number  353 


Barnes  and  Mitchell:  Kentriodon  from  California  3 


skull  lacking  the  extremity  of  the  rostrum,  the  occipital  shield, 
and  the  basicranium,  from  LACM  locality  1625,  and  31 
isolated  periotics  as  follows:  CAS  4348,  left  (holotype  of 
Grypolithax  pavida ),  CAS  4350,  left  (paratype  of  G.  pavida), 
both  from  CAS  locality  905;  LACM  21134,  right,  LACM 
449 1 6,  left,  LACM  63800,  left,  all  from  LACM  locality  1 625; 
LACM  2 1 238,  right,  and  LACM  48925,  left,  both  from  LACM 
locality  1557;  LACM  39696,  left,  from  LACM  locality  3232; 
LACM  41473,  left,  from  LACM  locality  1 655;  LACM  58893, 
left,  from  LACM  locality  6688;  LACM  75371,  left,  from 
LACM  locality  3208;  LACM  96150,  right,  LACM  121998, 
LACM  121999,  LACM  123477-123481,  eight  right,  LACM 
1 23482  and  1 23483  left,  all  from  LACM  locality  3 1 62;  LACM 
98712,  left,  from  LACM  locality  3160;  LACM  104094,  left, 
from  LACM  locality  1622;  LACM  123471-123473,  three 
right,  from  LACM  locality  4672;  LACM  123474,  left,  from 
LACM  locality  4874;  LACM  123475,  right,  and  LACM 
123476,  left,  from  LACM  locality  4956. 

FORMATION  AND  AGE.  The  holotype,  paratype,  and 
all  specimens  here  referred  to  Kentriodon  obscurus  were  col- 
lected from  several  localities  scattered  over  several  square 
kilometers  in  the  Sharktooth  Hill  Bonebed  in  the  upper  part 
of  the  Round  Mountain  Silt.  This  bonebed  is  a single  thin 
stratum  and  is  the  source  of  the  Sharktooth  Hill  Local  Fauna 
(Woodetal.,  1941;  Mitchell,  1 966:28-29;  Mitchell  and  Ted- 
ford,  1973;  Barnes,  1976).  It  is  correlated  with  the  Barstovian 
North  American  land  mammal  age,  the  “Temblor”  provi- 
sional provincial  molluscan  stage,  the  Relizian  or  Luisian 
foraminiferal  stage,  and  is  approximately  between  1 3 and  1 5 
million  years  old  (Wood  et  al.,  1941:31,  pi.  1 ; Weaver  et  al., 
1944:582,  pi.  1;  Evemden  et  al.,  1964;  Addicott,  1972;  Sav- 
age and  Barnes,  1972:133,  1 40;  Berggren  and  Van  Couvering, 
1974:  hg.  11;  Barnes,  1976:326-327;  1978:5-6;  Repennmg 
and  Tedford,  1977:  table  1).  Based  on  these  correlations,  the 
upper  part  of  the  Round  Mountain  Silt,  including  the  bone- 
bed, is  approximately  contemporaneous  with  the  Calvert 
Formation  in  Virginia  and  Maryland  (Gazin  and  Collins, 
1950:3;  Kellogg  and  Whitmore,  1957:1022;  Ray,  1976:  fig. 
1). 

DESCRIPTION  AND  COMPARISONS.  Skull.  The  re- 
ferred skull  (LACM  2 1 256,  Figs.  2-7)  of  Kentriodon  obscurus 
from  the  Sharktooth  Hill  Bonebed  is  incomplete  and  has 
suffered  pre-depositional  breakage  and  abrasion.  The  basi- 
cranium, occipital  area,  rostral  extremity,  vomer  and  ptery- 
goid sinus  fossae  have  been  broken  off.  The  sutures  between 
the  remaining  bones  were  unfused,  and  the  two  halves  were 
separated  along  the  midline  when  the  skull  was  found  in  the 
field.  We  have  re-assembled  it  in  its  presumed  original  con- 
figuration based  on  comparisons  with  specimens  of  Kentrio- 
don pernix  from  the  Calvert  Formation.  Due  to  some  dis- 
tortion, however,  the  left  supraorbital  process  fits  incorrectly 
and  attaches  too  low  on  the  skull  (Fig.  4a).  On  the  right  side 
(Fig.  4b)  the  supraorbital  process  is  in  the  correct  position. 
The  measurements  of  the  skull  are  as  follows:  total  length  as 
preserved,  1 93.5  mm;  breadth  of  rostrum  at  antorbital  notch- 
es, 67.7  mm;  breadth  of  cranium  at  antorbital  processes, 
122.3  mm;  interorbital  width,  1 15.2  mm. 

The  skull  is  nearly  the  same  size  as  both  the  holotype  and 


Kellogg’s  ( 1 927a)  referred  skull  of  K.  pernix.  All  three  appear 
to  represent  young  adult  animals  because  none  has  the  ad- 
vanced suture  fusion  and/or  extreme  development  of  rugos- 
ities and  processes  seen  on  skulls  of  extremely  old  individuals 
of  living  delphinoids.  Additionally,  in  the  holotype  skeleton 
of  K.  pernix  most  of  the  vertebral  epiphyses,  including  those 
on  cervicals  and  caudals,  are  tightly  appressed  to  the  vertebral 
centra  but  not  fused,  indicating  that  the  individual  had  not 
yet  achieved  physical  maturity.  Overall  skull  shape  and  pro- 
portions are  similar  in  the  two  species,  as  far  as  is  known.  It 
is  in  cranial  details  that  the  two  species  differ. 

The  open  mesorostral  gutter  on  the  distal  part  of  the  ros- 
trum is  apparently  characteristic  of  Kentriodon  pernix  (Kel- 
logg, 1927a:  pis.  2,  6),  but  is  uncharacteristic  of  species  in 
the  family  Kentriodontidae  as  a whole  (Barnes,  1978:  figs. 
14-17).  The  premaxillae  on  the  referred  skull  of  K.  obscurus 
are  partly  broken  away  distally,  but  in  addition  to  having  the 
usual  wide  open  mesorostral  gutter  at  the  proximal  end,  they 
begin  to  diverge  anteriorly  as  though  they  would  have  also 
left  the  gutter  exposed  distally  as  in  K.  pernix.  There  is  no 
indication  that  the  medial  margins  of  the  premaxillae  were 
elevated  adjacent  to  the  posterior  part  of  the  mesorostral 
gutter  as  they  are  in  K.  pernix.  The  anterior  (=  rostral)  parts 
of  the  premaxillae  are  comprised  of  bone  which  is  denser 
and  of  smoother  surface  than  the  adjacent  maxillae. 

On  the  skull  of  K.  obscurus,  the  premaxillary  foramina  are 
located  on  a transverse  line  between  the  antorbital  notches 
(Figs.  2-3).  These  foramina  are  located  more  posteriorly  on 
both  the  holotype  and  the  referred  skull  of  K.  pernix.  The 
premaxillary  sulci  associated  with  these  foramina  are  some- 
what damaged  from  breakage  and  abrasion,  but  between  the 
right  and  left  sides,  the  typical  odontocete  condition  of  three 
sulci  (anteromedial,  posteromedial,  and  posterolateral,  see 
Barnes,  1 978: 1 3;  Fordyce,  198 1 : 1034,  text-fig.  2)  can  be  seen. 
Medial  to  the  anteromedial  sulcus  and  the  premaxillary  fo- 
ramen, the  premaxillary  surface  is  rough  and  indicates  the 
area  where  the  nasal  plug  muscle  was  attached  (Lawrence 
and  Schevill,  1956:  fig.  23;  Mead,  1975).  The  part  of  the 
premaxilla  that  in  life  underlay  the  premaxillary  sac  (the 
premaxillary  sac  fossa  of  Fordyce,  1981:1035,  text-fig.  2; 
Mead,  1975)  lies  posterior  to  the  posteromedial  and  pos- 
terolateral sulci  and  lateral  to  the  nares.  This  area  is  also 
roughened,  but  this  is  the  result  of  postmortem  abrasion 
because  one  small  remnant  of  surficial  bone  on  the  left  pre- 
maxilla indicates  the  previous  existence  of  a thin,  smooth 
bone  surface  on  the  premaxillary  sac  fossa  as  is  typical  of 
delphinoids.  The  more  anterior  location  in  K.  obscurus,  com- 
pared with  K.  pernix,  of  the  premaxillary  foramina,  the  three 
associated  sulci,  and  concomitantly  the  anterior  margin  of 
the  premaxillary  sac  fossae,  is  primitive,  because  these  struc- 
tures have  moved  progressively  posteriorly  during  the  odon- 
tocete cranial  telescoping  process  (Miller,  1923).  The  area 
that  was  occupied  by  the  premaxillary  sacs  is  more  elongate 
anteroposteriorly  in  K.  obscurus,  and  we  consider  this  to  be 
primitive  as  well.  These  sacs  in  Kentriodon  might  have  been 
symmetrical  because  the  premaxillary  sac  fossae  are  not 
asymmetrical  as  they  are  in  modem  delphinids  (cf.  Mead, 
1975). 


4 Contributions  in  Science,  Number  353 


Barnes  and  Mitchell:  Kentriodon  from  California 


Both  species  of  Kentriodon  have  relatively  small  bony  nares 
that  are  narrowly  pointed  anteriorly.  This  is  also  typical  of 
many  other  species  of  Kentriodontidae  (see  Barnes,  1978: 
figs.  14-17),  and  contrasts  with  the  large,  round  nares  of 
species  of  Delphinidae. 

The  lateral  margin  of  the  maxilla  dorsal  to  the  posterior 
end  of  the  tooth  row  and  medial  to  the  antorbital  notch 
typically  is  formed  into  a relatively  thick,  horizontally  pro- 
jecting shelf  in  most  kentriodontids.  In  K.  obscurus,  this  part 
of  the  maxilla  is  proportionally  wider  and  has  a more  squared 
off  and  vertical  lateral  margin  than  in  K.  pernix,  in  which 
the  maxilla  has  a thinner,  more  rounded  margin.  The  ant- 
orbital notches  of  K.  obscurus  are  relatively  narrower  than 
those  of  K.  pernix,  because  in  K.  obscurus,  both  the  lacrimal 
and  the  maxilla  project  in  a more  anteromedial  direction, 
thereby  giving  the  antorbital  process  a different  shape  (Fig. 

13). 

On  the  supraorbital  process,  both  the  frontal  and  maxilla 
are  significantly  thicker  and  thus  more  convex  or  domed  in 
K.  obscurus  than  in  K.  pernix.  The  lateral  margins  of  the 
maxillae  have  been  both  chipped  and  abraded  over  the  orbit 
on  the  skull  of  K.  obscurus.  The  frontal  is  therefore  probably 
exposed  more  in  dorsal  view  than  it  was  in  life. 

The  postorbital  process  of  the  frontal  of  K.  obscurus  is 
narrower,  shorter,  and  more  tapered  distally  than  in  K.  per- 
nix. These  differences  do  not  appear  to  be  related  to  ontogeny 
because  the  juvenile  skull  that  we  refer  to  K.  pernix  (USNM 
21027)  from  the  Calvert  Formation  is  smaller  than  the  skull 
of  K.  obscurus,  has  characters  indicating  physical  immatu- 
rity, but  has  substantially  longer  postorbital  processes.  Ad- 
ditionally, the  postorbital  processes  on  the  three  skulls  of 
Kentriodon  pernix  from  the  Calvert  Formation  have  a hook- 
like shape,  in  contrast  to  the  straight  process  of  K.  obscurus. 

Among  modem  Delphinidae,  there  is  a fossa  located  at  the 
posterolateral  comer  of  the  palate  lateral  to  the  pterygoid 
sinus  fossa  and  which  marks  the  position  of  an  air  sinus  called 
the  anterior  sinus  (see  Fraser  and  Purves,  1960).  This  area 
of  the  palate  of  K.  obscurus  is  convex  and  shows  no  devel- 
opment of  such  a fossa.  On  skulls  of  K.  pernix,  this  area  of 
the  palate  is  distinctly  less  convex  so  that  a transverse  section 
through  the  proximal  part  of  the  rostrum  is  nearly  V-shaped. 
Neither  species  has  the  type  of  fossa  or  concave  area  that  in 
some  living  odontocetes  marks  the  location  of  an  anterior 
sinus,  and  there  probably  was  no  such  sinus  present  in  either 
species  of  Kentriodon. 

The  fossa  for  the  lobe  of  the  pterygoid  air  sinus  that  filled 
the  pterygoid  hamulus  is  larger  and  extends  farther  anteriorly 
in  K.  obscurus  than  in  K.  pernix.  The  roof  of  this  pterygoid 
sinus  fossa  is  marked  by  a tapered  cavity  on  the  ventral 
surface  of  the  left  palatine  bone.  The  palatines  concomitantly 
extend  farther  anteriorly  and  are  more  pointed  in  K.  obscurus. 
The  anterior-most  extent  of  the  palatines  in  K.  obscurus  is 
35  mm  anterior  to  the  antorbital  notch,  whereas  the  same 
parameter  on  the  holotype  of  K.  pernix  is  only  10  mm  and 
on  Kellogg’s  referred  skull  (USNM  10670)  it  is  14.4  mm. 

As  on  the  referred  skull  (USNM  10670)  of  K.  pernix,  the 
ventral  surface  of  the  vomer  was  only  exposed  in  a very 
narrow  opening  between  the  maxillae  at  about  the  middle  of 


the  rostrum  of  K.  obscurus  (Figs.  14a,  b).  In  both  species, 
posterior  palatine  foramina  flank  this  area,  and  slightly  far- 
ther anteriorly  a single  anterior  palatine  foramen  occurs  in 
each  maxilla  adjacent  to  the  posterior-most  palatal  exposure 
of  the  premaxillae. 

The  teeth  of  K.  obscurus  were  significantly  smaller  in  di- 
ameter than  those  of  K.  pernix  (Figs.  14a,  b).  In  K.  pernix, 
throughout  the  tooth  row,  the  alveoli  have  uniform  diameter 
(3  mm)  and  nearly  equal  spacing.  In  K.  obscurus,  however, 
the  alveoli  that  are  still  intact  on  the  right  side  located  just 
posterior  to  the  mid-length  of  the  rostrum  are  only  1.5  mm 
in  diameter;  one-half  the  size  of  those  in  K.  pernix.  The 
alveolar  rows  on  both  sides  of  the  specimen  of  K.  obscurus 
are  incomplete  because  of  breakage  and  abrasion.  A row  of 
eight  consecutive  alveoli  in  a distance  of  32  mm  in  the  right 
maxilla  indicates  that  the  species  had  more  teeth  than  K. 
pernix,  which  Kellogg  (1927a:32)  estimated  at  about  40  on 
each  side  of  each  upper  jaw.  The  alveoli  of  K.  obscurus  are 
oriented  in  the  maxilla  so  that  they  face  more  laterally  than 
in  K.  pernix.  The  alveolar  rows  do  not  extend  as  far  poste- 
riorly on  the  palate,  and  the  posterior  end  of  each  tooth  row 
curves  medially  toward  the  midline. 

Posterior  to  the  orbit,  K.  obscurus  has  a large,  oval  fossa 
in  the  ventral  side  of  the  supraorbital  process  of  the  frontal 
that  measures  approximately  15  by  20  mm.  In  modem  del- 
phinoids,  a fossa  in  this  location  holds  the  postorbital  lobe 
of  the  pterygoid  air  sinus  (Fraser  and  Purves,  1960).  We 
conclude  that  K.  obscurus  had  a relatively  large  sinus  here. 
On  both  the  holotype  and  Kellogg’s  referred  skull  (USNM 
10670)  of  K.  pernix,  there  is  only  a slight  depression  at  this 
place.  This  apparent  difference  in  air  sinus  size  is  not  clearly 
diagnostic  because  the  juvenile  referred  skull,  USNM  21027 
of  K.  pernix,  has  a deep  recess  for  the  sinus  at  this  place. 
Without  a larger  sample  of  specimens  we  cannot  determine 
whether  the  size  or  the  extent  of  invasion  of  bone  by  the 
sinus  is  variable  in  K.  pernix,  whether  it  is  larger  in  geolog- 
ically more  recent  individuals  of  K.  pernix  that  occur  higher 
in  the  Calvert  Formation,  or  whether  it  is  a significant  taxo- 
nomic character  that  separates  species. 

Periotic.  When  Kellogg  (1931)  described  Grypolithax  ob- 
scura  and  G.  pavida  from  the  Sharktooth  Hill  Bonebed,  he 
noted  only  a few  differences  between  their  holotypes.  He 
noted  that  the  periotics  of  G.  pavida  had  a flatter  cerebral 
surface.  We  now  benefit  from  having  a larger  sample  to  study 
and  find  that,  within  the  anticipated  range  of  morphology  of 
what  we  interpret  as  one  species,  the  presence  of  a flat  cerebral 
surface  is  variable. 

Among  the  31  isolated  periotics  from  the  Sharktooth  Hill 
Bonebed  that  we  refer  to  Kentriodon  obscurus,  there  are  at 
least  six  (LACM63800, 48925, 21238, 4 1473, 7537 1,1 23476) 
that  are  nearly  identical  in  morphology  to  the  periotic  of  the 
holotype  of  K.  pernix  (USNM  8060).  Each  of  the  others  has 
some  minor  degree  of  difference.  There  was  no  periotic  found 
with  the  skull  we  have  referred  to  K.  obscurus,  and  except 
for  the  holotype  of  K.  pernix,  there  has  been  found  only  one 
other  skull  of  Kentriodon  sp.  in  the  Calvert  Formation  that 
has  an  associated  periotic.  This  specimen,  USNM  187313, 
which  we  believe  is  a different  and  probably  undescribed 


Contributions  in  Science,  Number  353 


Barnes  and  Mitchell:  Kentriodon  from  California  5 


6 Contributions  in  Science,  Number  353 


Barnes  and  Mitchell:  Kentriodon  from  California 


Figure  2.  Kentriodon  obscurus  (Kellogg,  1931),  referred  skull,  LACM  21256  from  LACM  locality  1625,  dorsal  view,  natural  size. 


fmx 


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Contributions  in  Science,  Number  353 


Barnes  and  Mitchell:  Kentriodon  from  California  7 


Figure  3.  Kentriodon  obscurus  (Kellogg,  1931),  referred  skull,  LACM  21256  from  LACM  locality  1625.  dorsal  view,  natural  size.  Abbreviations:  aon  — antorbital  notch,  fmx 
maxillary  foramen,  fpmx— premaxillary  foramen,  Fr— frontal.  La— lacrimal,  mrg— mesorostral  gutter,  Mx— maxilla,  n — nanal  opening,  Pmx  — premaxilla. 


8 Contributions  in  Science,  Number  353 


Barnes  and  Mitchell:  Kentriodon  from  California 


Kentriodon  obscurus  (Kellogg,  1931),  referred  skull,  LACM  21256  from  LACM  locality  1625,  a,  left  lateral  view;  b,  right  lateral  view,  natural  size. 


I 


Contributions  in  Science,  Number  353 


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Table  1.  Some  characters  differentiating  Kentriodon  pernix  and  K. 
obscurus.  An  asterisk  denotes  the  more  derived  character  states,  as 
deduced  from  the  morphology  of  Agorophius  and  species  of  Squal- 
odontidae. 

Kentriodon  obscurus  Kentriodon  pernix 


1.  Premaxillary  foramina  be- 
tween level  of  antorbital 
notches. 

2.  Lateral  maxillary  margin 
anterior  to  antorbital 
notches  thick  and  squared 
off.* 

3.  Antorbital  notch  narrow. 

4.  Premaxillae  flat  in  area  of 
attachment  of  nasal  plug 
muscles.* 

5.  Postorbital  process  of 
frontal  short  and  thick.* 

6.  Alveoli  for  teeth  small 
(circa  1.5  mm  in  diame- 
ter). 

7.  Posterior  end  of  alveolar 
row  bends  medially  on 
palate. 

8.  Teeth  implanted  in  mid- 
dle and  posterior  part  of 
alveolar  row  so  as  to  face 
ventrolaterally.* 

9.  Palatines  and  pterygoid 
air  sinus  fossae  on  palate 
extend  farther  anteriorly 
beyond  location  of  antor- 
bital processes.* 

10.  Posterior  part  of  palate 
lateral  to  fossae  for  ptery- 
goid air  sinus  convex. 

1 1.  Large  fossa  in  ventral  sur- 
face of  frontal  for  postor- 
bital lobe  of  pterygoid  air 
sinus.* 

12.  Internal  acoustic  meatus 
of  periotic  elliptical  in 
shape. 


1 . Foramina  posterior  to 
level  of  notches.* 

2.  Lateral  margin  thin  and 
rounded. 

3.  Antorbital  notch  wider.* 

4.  Premaxillae  elevated  me- 
dially. 

5.  Process  long  and  curved. 

6.  Alveoli  larger  (circa  3 mm 
in  diameter).* 

7.  Posterior  end  of  alveolar 
row  follows  lateral  margin 
of  palate.* 

8.  Teeth  face  more  ventrally. 


9.  Palatines  and  fossae  do 
not  extend  beyond  level 
of  antorbital  processes. 


10.  Palatal  surface  less  con- 


1 1.  Fossa  small  or  absent. 


12.  Meatus  circular  in  shape.* 


species  of  Kentriodon  was  collected  from  Zone  14,  near  the 
top  of  the  Calvert  Formation,  and  the  skull,  periotic,  and 
postcranial  skeleton  show  several  significant  differences  from 
the  holotype  and  referred  specimen  of  K.  pernix.  The  periotic 
of  this  specimen,  USNM  1 8731 3 (Figs.  8a,  9a),  has  a prom- 
inent, elevated,  flat  area  on  its  cerebral  surface  that  is  more 
pronounced  than  on  any  of  the  other  periotics  referred  to 


Barnes  and  Mitchell:  Kentriodon  from  California  9 


10  Contributions  in  Science,  Number  353 


Barnes  and  Mitchell:  Kentriodon  from  California 


I 


Figure  6.  Kentriodon  obscurus  (Kellogg,  1931),  referred  skull,  LACM  21256  from  LACM  1625,  ventral  view,  natural  size. 


sod 


Contributions  in  Science,  Number  353 


Barnes  and  Mitchell:  Kentriodon  from  California  1 1 


Figure  7.  Kentriodon  obscurus  (Kellogg,  1931),  referred  skull,  LACM  21256  from  LACM  locality  1625,  ventral  view,  natural  size.  Abbreviations:  aon-antorbital  notch,  fap— 
anterior  palatine  foramen,  fio— orbital  apertures  of  the  infraorbital  foramen.  Fr— frontal.  La— lacrimal,  Mx  — maxilla,  n — nanal  opening,  pos— fossa  for  postorbital  lobe  of  the 
pterygoid  sinus  of  the  middle  ear  air  sinus  system,  Pmx— premaxilla,  pts  — fossa  for  pterygoid  sinus  of  the  middle  ear  air  sinus  system,  Vo— cleft  that  originally  contained  the 
palatal  exposure  of  the  vomer. 


Figure  8.  Periotics  of  Kentriodon  Kellogg,  1 927:  a,  Kentriodon  sp.,  Calvert  Fm.,  Virginia,  USNM  187313,  right;  b,  Kentriodon  pernix  Kellogg, 
1927,  holotype,  USNM  8060,  left;  c,  Kentriodon  sp.,  Calvert  Fm.,  Virginia,  USNM  214754,  right;  Kentriodon  obscurus  (Kellogg,  1931)  from 
the  Sharktooth  Hill  Bonebed;  d,  LACM  41473,  left;  e,  CAS  4349,  holotype,  right;  f,  CAS  4348,  left  (holotype  of  Grypolithax pavida  Kellogg, 
1931);  g,  CAS  4350,  left  (paratype  of  Grypolithax  pavida)-,  h,  LACM  75371,  right;  i,  LACM  63800,  left;  j,  LACM  48925,  left;  k,  CAS  4347,  i 

paratype,  left;  1,  LACM  21238,  right;  m,  LACM  121998,  right;  n,  LACM  58893,  left;  o,  LACM  104094,  left;  p,  LACM  44916,  left;  q,  LACM 
98712,  left;  r,  LACM  21 134,  right;  s,  LACM  39696,  left;  t,  LACM  96150,  right;  all  figures  are  stereophotographs  of  the  cerebral  (or  dorsal) 
surface,  natural  size. 

i 

l 

( 


Kentriodon.  Such  a flat  surface  is  present,  however,  in  varying 
lesser  degrees  on  eight  of  the  periotics  from  the  Sharktooth 
Hill  Bonebed  referred  to  K.  obscurus  (including  the  holotype 
of  Grypolithax  pavida ),  as  well  as  on  the  holotype  periotic 
of  K.  pernix.  Another  isolated  periotic  (USNM  214754,  Figs. 
8c,  9c)  from  the  Calvert  Formation  in  Virginia  closely  match- 
es the  holotype  periotic  of  K.  pernix,  and  although  it  has  a 
more  rounded  cerebral  surface,  we  refer  it  to  that  species. 

All  of  the  periotics  from  the  Sharktooth  Hill  Bonebed  that 
we  refer  to  Kentriodon  obscurus  resemble  the  holotype  peri- 


otic of  K.  pernix  by  having  the  following  characters:  (1)  sim- 
ilar size;  (2)  relatively  small  anterior  and  posterior  processes; 

(3)  an  overall  sinuosity  in  either  cerebral  or  ventral  view 
owing  to  the  fact  that  the  anterior  process  is  bent  medially 
and  the  posterior  process  is  bent  laterally;  (4)  cochlear  portion 
relatively  small  and  broadly  joined  to  the  body  of  the  periotic, 
not  narrowly  joined  to  the  body  and  extended  medially  as 
in  many  primitive  odontocetes;  (5)  posterior  process  bent  | 
ventrally  at  a sharp  angle  from  the  body  of  the  periotic  there- 
by forming  a sharp  peak  or  angle  on  the  posterior  part  of  the 


12  Contributions  in  Science,  Number  353 


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Contributions  in  Science,  Number  353 


Barnes  and  Mitcheil:  Kentriodon  from  California  13 


Figure  9.  Periotics  of  Kentriodon  Kellogg,  1 927:  a,  Kentriodon  sp.,  Calvert  Fm.,  Virginia,  USNM  187313,  right;  b,  Kentriodon  pernix  Kellogg, 
1927,  holotype,  USNM  8060,  left;  c,  Kentriodon  sp.,  Calvert  Fm.,  Virginia,  USNM  214754,  right;  Kentriodon  obscurus  (Kellogg,  1931)  from 
the  Sharktooth  Hill  Bonebed;  d,  LACM  41473,  left;  e,  CAS  4349,  holotype,  right;  f,  CAS  4348,  left  (holotype  of  Grypolithax  pavida  Kellogg, 
1931);  g,  CAS  4350,  left  (paratype  of  Grypolithax  pavida);  h,  LACM  75371,  right;  i,  LACM  63800,  left;  j,  LACM  48925,  left;  k,  CAS  4347, 
paratype,  left;  1,  LACM  21238,  right;  m,  LACM  121998,  right;  n,  LACM  58893,  left;  o,  LACM  104094,  left;  p,  LACM  44916,  left;  q,  LACM 
98712,  left;  r,  LACM  21134,  right;  s,  LACM  39696,  left;  t,  LACM  96150,  right;  all  figures  are  stereophotographs  of  the  tympanic  (or  ventral) 
surface,  natural  size. 


cerebral  surface;  (6)  anterior  process  bent  anteroventrally 
and  having  a groove  or  pit  on  its  medial  side;  (7)  the  cleft 
between  the  anterior  process  and  the  cochlear  portion  bearing 
a small  crease  between  the  groove  for  the  tensor  tympani 
muscle  and  the  cochlear  portion;  (8)  articular  facet  for  the 
auditory  bulla  on  the  posterior  process  large,  concave  and 
striated;  and  (9)  a raised  rugosity  lateral  to  the  fossa  for  the 
head  of  the  malleus. 

Statistical  analysis  of  the  Sharktooth  Hill  Bonebed  sample 


of  K.  obscurus  periotics  (Table  2)  proved  useful.  The  param- 
eters chosen  were  width  and  length  measurements.  The  mea- 
surement of  the  cochlear  portion  is  probably  less  variable 
individually  or  allometrically  than  the  other  two,  because  the 
sizes  of  the  anterior  and  posterior  processes  apparently  change 
during  ontogeny.  Measurements  of  the  holotype  of  K.  ob- 
scurus fall  within  the  range  of  measurements  for  the  sample, 
and  they  differ  from  the  mean  less  than  the  standard  devia- 
tion in  all  three  parameters.  The  same  situation  exists  when 


14  Contributions  in  Science,  Number  353 


Barnes  and  Mitchell:  Kentriodon  from  California 


Contributions  in  Science,  Number  353 


Barnes  and  Mitchell:  Kentriodon  from  California  15 


Table  2.  Statistical  comparisons  of  the  holotype  periotic  of  Kentriodon  pernix  and  the  holotype  and  referred  periotics  of  K.  obscurus  from  the 
Sharktooth  Hill  Bonebed. 


Bonebed  sample  of  K.  obscurus* 

K.  obscurus  holotype 

K.  pernix  holotype 

Measurement 

Number 

Observed 

range 

Mean 

Standard 

deviation 

Coefficient 
of  variation 

Measure- 

ment 

Difference 
from  mean 

Measure- 

ment 

Difference 
from  mean 

Total  length 

27 

25.2-29.4 

27.2 

1.2 

4.41 

26.9 

-0.3 

28.3 

1.1 

Width  at  pars 
cochlearis 

29 

16.5-19.5 

18.0 

0.8 

4.44 

18.2 

0.2 

17.4 

-0.6 

Antero-posterior 
dimension  of 
pars  cochlearis 

28 

13.2-15.9 

14.7 

0.7 

4.76 

14.1 

-0.6 

14.2 

-0.5 

* Sample  includes  paratype  of  K.  obscurus  and  holotype  and  paratype  of  Grypolithax  pavida. 


the  holotype  of  K.  pernix  is  compared  to  the  sample  of  K. 
obscurus,  except  that  it  differs  somewhat  more  from  the  mean 
than  does  the  holotype  of  K.  obscurus,  but  is  still  within  the 
standard  deviation. 

The  similarities  among  these  periotics  from  the  Atlantic 
and  Pacific  coasts  are  not  surprising.  Kasuya  (1973:72)  and 
Barnes  (1976:321-322,  327)  have  pointed  out  that  periotics 
of  congeneric  species  of  cetaceans  are  very  similar  and  that 
in  some  instances  closely  related  modem  or  fossil  species 
simply  may  not  be  differentiated  based  on  periotics  alone. 

Among  the  periotics  of  K.  obscurus,  some  characters  are 
variable.  These  are:  ( 1 ) shape  and  relative  size  of  the  posterior 
articular  facet  for  the  bulla;  (2)  degree  of  rugosity  of  this 
articular  facet;  (3)  position  of  the  groove  or  pit  on  the  medial 
side  of  the  anterior  process;  (4)  presence  or  absence  of  a 
tuberosity  on  the  posterior  side  of  the  cochlear  portion  be- 
tween the  fenestra  rotunda  and  the  orifice  of  the  aquaeductus 
cochleae;  (5)  extent  of  development  of  an  attenuated  groove 
in  the  anterior  margin  of  the  internal  acoustic  meatus  at  the 
cerebral  orifice  of  aquaeductus  Fal/opii;  and  (6)  extent  of 
development  of  a flat  area  on  the  cerebral  surface  of  the  body 
of  the  periotic  in  contrast  with  a rounded,  convex  surface. 
The  only  character  that  we  recognize  whereby  the  holotype 
periotic  of  Kentriodon  pernix  may  be  separated  from  this 
sample  of  periotics  of  K.  obscurus  is  its  more  circular  internal 
acoustic  meatus.  Every  other  character  exists  in  at  least  one 
of  the  periotics  from  the  Sharktooth  Hill  Bonebed  that  we 
refer  to  K.  obscurus. 

The  only  other  significantly  large,  documented  sample  of 
periotics  of  a species  of  Kentriodontidae  is  the  series  of  Lioli- 
thax  kernensis  from  the  Sharktooth  Hill  Bonebed  that  was 
described  and  analyzed  by  Bames  (1978).  The  periotics  from 
the  bonebed  that  we  now  refer  to  Kentriodon  obscurus  are 
as  variable  as  the  sample  Bames  referred  to  L.  kernensis. 
These  periotics  are  not  referable  to  any  of  the  other  odon- 
tocetes  that  Kellogg  (193 1)  named  based  upon  periotics  from 
the  Sharktooth  Hill  Bonebed. 

Among  these  other  species,  the  problematic  odontocete 
Platylithax  robusta  Kellogg,  1931,  the  primitive  sperm  whale 
Aulophyseter  morricei  Kellogg,  1927,  the  platanistoid  dol- 
phin “ Squalodon"  errabundus  Kellogg,  1931,  and  the  ken- 


triodontid  dolphin  Liolithax  kernensis  all  have  periotics  that 
are  substantially  different  from  those  belonging  to  Kentri- 
odon. Periotics  of  another  dolphin,  Loxolithax  sinuosa  Kel- 
logg, 1931,  differ  subtly  by  being  flatter  dorsoventrally,  hav- 
ing a larger  cochlear  portion,  a differently  shaped  posterior 
process,  a larger  fossa  for  the  head  of  the  malleus,  a groove 
on  the  lateral  surface  of  the  anterior  process,  and  by  lacking 
any  notable  flattening  of  the  cerebral  surface.  Periotics  of  the 
two  species  of  Lamprolithax  Kellogg,  1931,  differ  from  those 
of  Kentriodon  in  many  of  the  same  ways  as  do  those  of 
Loxolithax  sinuosa.  Nannolithax  gracilis  Kellogg,  1931,  has 
a smaller  periotic  with  a large  anterior  process.  Oedolithax 
mira  Kellogg,  1931,  has  a larger  periotic  with  a deeper  cleft 
separating  the  anterior  process  from  the  cochlear  portion. 

cf.  Kentriodon  sp. 

Figures  10-11 

cf.  Kentriodon  Kellogg,  1927.  Bames,  1976:326. 

REFERRED  SPECIMENS.  LACM  29549,  left  periotic, 
collected  by  Richard  W.  Huddleston  in  1969  from  LACM 
locality  3066,  Kem  River,  Kem  County,  California;  and 
UCMP  83605,  right  periotic,  collected  by  Richard  C.  Bishop 
from  UCMP  locality  V-6953,  Kem  County,  California. 

FORMATION  AND  AGE.  Both  of  these  specimens  are 
from  localities  stratigraphically  below  the  Sharktooth  Hill 
Bonebed  in  the  lower  part  of  the  Round  Mountain  Silt.  This 
part  of  the  formation  is  early  Middle  Miocene  in  age,  about 
15  to  19  million  years  old  (see  Bames,  1976:326).  Savage 
and  Bames  (1972:133)  have  reported  Hemingfordian  land 
mammals  from  these  same  strata.  Addicott  (1972)  charac- 
terized the  age  of  these  beds  further  as  representing  part  of 
the  “Temblor”  provisional  provincial  molluscan  stage  and 
the  upper  part  of  the  Saucesian  or  the  Relizian  foraminiferal 
stage. 

DESCRIPTION  AND  COMPARISONS.  These  two 
periotics  differ  from  each  other  in  both  size  and  morphology 
and  may  actually  represent  two  different  species.  Each  shares 
characters  in  common  with  both  Kentriodon  obscurus  and 
K.  pernix,  such  as  the  strong  apex  on  the  cerebral  surface 


16  Contributions  in  Science,  Number  353 


Barnes  and  Mitchell:  Kentriodon  from  California 


Figure  10.  cf.  Kentriodon  sp.,  left  periotic,  LACM  29549  from 
LACM  locality  3066;  a,  cerebral  or  dorsal  view;  b,  tympanic  or 
ventral  view;  both  natural  size. 

posterolateral  to  the  orifice  of  aquaeductus  vestibu/i,  small 
internal  acoustic  meatus,  and  anteromedially  projecting  an- 
terior process— all  characters  that  could  be  regarded  as  prim- 
itive. The  smaller  of  the  two  periotics,  UCMP  83605,  has 
an  anteroposteriorly  elongate  cochlear  portion  as  in  other 
specimens  of  Kentriodon.  The  larger  periotic,  LACM  29549, 
has  a relatively  smaller  and  more  globose  cochlear  portion, 
and  in  this  respect  it  resembles  the  holotype  periotic  of  the 
primitive  kentriodontid  Kampholophos  serrulus  Rensberger, 
1969  (see  Rensberger,  1969:  pi.  4,  figs.  f-h). 

aff.  Delphinodon  dividum  True,  1912 

Figure  12 

cf.  Delphinodon  dividum  True,  1912.  Barnes,  1976:326. 

REFERRED  SPECIMEN.  LACM  41041,  left  periotic  with 
stapes,  lacking  extremity  of  posterior  process,  collected  by 
the  late  John  E.  Fitch,  about  1970  from  LACM  locality  6602, 
“Barker’s  Ranch  Faunal  Site,”  Kern  County,  California. 

FORMATION  AND  AGE.  This  specimen  is  from  a lo- 
cality in  the  lower  part  of  the  Round  Mountain  Silt,  below 
the  Sharktooth  Hill  Bonebed,  in  roughly  the  same  strata  as 
the  two  previously  described  periotics.  Its  age  is  likewise  early 
Middle  Miocene  and  between  approximately  1 5 and  19  mil- 
lion years  old. 

DESCRIPTION  AND  COMPARISONS.  The  compari- 
sons that  we  have  made  are  specifically  only  with  Delphi- 
nodon dividum.  because  that  is  the  only  species  of  the  genus 
for  which  a periotic  is  known.  The  holotype  is  a specimen 
collected  from  the  Calvert  Formation  and  includes  a skull, 
mandible,  and  part  of  the  postcranial  skeleton.  The  type 
species  of  the  genus,  D.  mento  Cope,  1868,  as  fixed  by  Hay 
(1902),  is  known  only  by  teeth. 

As  noted  by  Barnes  (1976:326)  this  periotic  closely  resem- 
bles that  of  Delphinodon  dividum.  Kentriodon  spp.  and  Del- 
phinodon dividum  are  closely  related  (cf.  Kellogg,  1927a,  1928: 
67-69;  Slijper,  1936:556),  and  both  have  been  classified  in 
the  subfamily  Kentriodontinae  (Barnes,  1978).  The  periotic 
with  the  holotype  of  Delphinodon  dividum  (see  True,  1912; 
pi.  25,  figs.  6,  7)  has  many  characters  in  common  with  the 
isolated  periotic  (LACM  41041)  from  Kern  County.  Notable 
among  these  are  the  large  fossa  for  the  head  of  the  malleus, 
the  globose  anterior  process  which  is  bent  medially,  and  the 
small,  spherical  cochlear  portion.  The  periotic  of  D.  dividum 
is  somewhat  similar  in  overall  shape  to  periotics  of  Kentri- 
odon obscurus  and  K.  pernix,  but  differs  by  not  having  the 


Figure  11.  cf.  Kentriodon  sp.,  right  periotic,  UCMP  83605  from 
UCMP  locality  V-6953;  a,  cerebral  or  dorsal  view;  b,  tympanic  or 
ventral  view;  c,  lateral  view;  all  natural  size. 


cochlear  portion  as  broad  anteroposteriorly,  by  having  the 
anterior  process  bent  more  medially,  and  by  having  the  coch- 
lear portion  separated  from  the  anterior  process  by  a much 
deeper  fissure.  The  isolated  periotic  from  Kern  County 
(LACM  41041)  differs  from  the  holotype  of  D.  dividum  by 
being  smaller,  having  a relatively  smaller  posterior  articular 
facet  for  the  bulla,  and  by  having  a relatively  smaller  and 
narrower  internal  acoustic  meatus.  The  latter  character  may 
be  primitive  in  comparison  with  D.  dividum.  We  believe  this 
isolated  periotic  from  the  Round  Mountain  Silt  is  congeneric 
with  Delphinodon  dividum  and  represents  an  earlier,  more 
primitive  species. 

A very  closely  related,  if  not  identical,  species  is  repre- 
sented by  a periotic  from  correlative  rocks  in  Japan.  Okazaki 
(1976:37-38,  text-fig.  6,  pi.  11,  figs,  la-c  (where  the  scale  is 
incorrect))  has  identified  that  periotic,  collected  from  the  late 
Early  or  early  Middle  Miocene  Nataki  Formation,  as  a rhab- 
dosteid  dolphin,  Eurhinodelphis  sp.  B.  We  do  not  concur 
with  that  generic  allocation  because  the  periotic  described 
by  Okazaki  does  not  closely  resemble  periotics  found  in  skulls 
of  Eurhinodelphis  spp.  collected  from  the  Calvert  Formation 
in  Maryland  and  Virginia  (USNM  collections).  Instead,  we 
believe  the  periotic  from  Japan  very  closely  resembles  the 
periotic  of  Delphinodon  dividum,  and  the  one  from  California 
that  we  identify  here  as  aff.  Delphinodon  dividum,  although 
it  apparently  is  considerably  larger  than  the  latter.  We  con- 
clude that  the  specimen  from  the  Nataki  Formation  should 


a b 

Figure  1 2.  aff.  Delphinodon  dividum  T rue,  1912,  left  periotic,  LACM 
41041  from  LACM  locality  6602;  a,  cerebral  or  dorsal  view;  b, 
tympanic  or  ventral  view;  both  natural  size. 


Contributions  in  Science,  Number  353 


Barnes  and  Mitchell:  Kentriodon  from  California  17 


Figure  13.  Reconstructions  of  dorsal  views  of  skulls  of  Kentriodon  Kellogg,  1927;  a,  Kentriodon  pernix  Kellogg,  1927,  based  on  the  holotype, 
USNM  8060,  and  the  referred  specimen,  USNM  10670;  b,  Kentriodon  obscurus  (Kellogg,  1931),  based  on  the  referred  specimen,  LACM 
21256,  with  outline  of  the  rostral  extremity  and  brain  case  from  K.  pernix;  both  at  different  scales,  but  reduced  to  the  same  brain  case  length 
(antorbital  notches  to  condyles),  (a  from  Kellogg,  1927:  pis.  2,  6,  and  Barnes,  1978:  fig.  14,  c;  b from  our  Fig.  2.) 


18  Contributions  in  Science,  Number  353 


Barnes  and  Mitchell:  Kentriodon  from  California 


Figure  14.  Reconstructions  of  ventral  views  of  skulls  of  Kentriodon  Kellogg,  1927;  a,  Kentriodon  pernix  Kellogg,  1927,  based  on  the  holotype, 
USNM  8060,  and  the  referred  specimen,  USNM  10670,  with  the  dentition  omitted  on  one  side  so  that  the  size  and  number  of  alveoli  may 
be  seen;  b,  Kentriodon  obscurus  (Kellogg,  1931),  based  on  the  referred  specimen  LACM  21256,  with  outline  of  the  rostral  extremity  and  brain 
case  from  K.  pernix;  both  at  different  scales,  but  reduced  to  the  same  brain  case  length,  (a  from  Kellogg,  1927:  pis.  4,  5,  7 and  8;  b from  our 
Fig.  6.) 


Contributions  in  Science,  Number  353 


Barnes  and  Mitchell:  Kentriodon  from  California  19 


be  re-identified  as  Kentriodontinae,  aff.  Delphinodon  divi- 
dum. 

DISCUSSION 

RELATIVE  ABUNDANCE  OF 
KENT RIODON  OBSCURUS  IN  THE 
SHARKTOOTH  HILL  BONEBED 

A list  of  1 5 odontocetes  in  the  Sharktooth  Hill  Bonebed  given 
by  Barnes  ( 1976:327,  table  3)  included  1 1 named  species  and 
four  possibly  undescribed  species.  Grypolithax  pavida  may 
now  be  deleted  from  that  list  because  it  is  a synonym  of  K. 
obscurus.  Each  of  the  14  remaining  species  is  recognized  by 
periotics.  Some  of  the  species  in  polytypic  genera  that  are 
known  only  by  periotics  may  be  questionably  valid  (Barnes, 
1976:327).  The  abundant  sperm  whale,  Aulophyseter  mor- 
ricei,  with  quite  distinctive  periotics,  is  known  by  other  skel- 
etal elements.  The  platanistoid  “ Squalodon"  errabundus  also 
has  distinctive  periotics  and  is  known  from  other  skeletal 
elements,  however,  it  is  relatively  rare  in  the  bonebed,  and 
we  (unpublished  data)  have  been  able  to  collect  or  locate  only 
31  of  the  highly  unique  periotics  of  that  species.  Similarly, 
the  distinctive  periotics  of  the  kentriodontid  Liolithax  ker- 
nensis  total  only  42  in  number  (Barnes,  1978).  Only  Platy- 
lithax  robusta.  known  solely  by  the  holotype,  and  the  other 
four  unidentified  odontocete  species  are  rarer  in  samples  than 
Kentriodon  obscurus.  The  total  sample  of  K.  obscurus  amounts 
to  only  one  skull  and  31  periotics,  and  on  this  evidence  the 
species  ranks  as  one  of  the  rarest  odontocetes  in  the  Shark- 
tooth Hill  Bonebed.  The  remaining  hundreds  of  periotics  in 
museum  collections  can  be  identified  as  belonging  to  species 
of  the  genera  Loxolithax  Kellogg,  1931,  Oedolithax  Kellogg, 
1931,  Lamprolithax,  or  Nannolithax  Kellogg,  1931. 

ZOOGEOGRAPHY 

Fossil  Kentriodontidae  have  been  reported  from  the  North 
Atlantic,  the  South  and  North  Pacific,  and  the  Paratethys 
regions.  All  the  species  included  by  Barnes  (1978:26)  in  this 
family  were  Middle  or  Late  Miocene  in  age.  Fordyce  (1980: 
328)  has  subsequently  reported  a kentriodontid  from  Late 
Oligocene  rocks  of  New  Zealand.  The  fossils  from  early  Mid- 
dle Miocene  rocks  in  California  identified  as  cf.  Kentriodon 
sp.  and  aff.  Delphinodon  dividum  in  this  paper  constitute  the 
earliest  records  for  the  family  in  the  North  Pacific  region. 
These,  along  with  the  sample  we  regard  as  Kentriodon  ob- 
scurus from  the  Sharktooth  Hill  Bonebed,  the  kentriodontids 
from  Japan,  and  Late  Miocene  species  from  California 
(Barnes,  1976)  suggest  that  a considerable  diversity  of  species 
in  this  family  inhabited  the  North  Pacific  Ocean  during  the 
Miocene.  This  diversity  equals  that  known  for  kentriodontids 
during  the  Middle  and  Late  Miocene  in  the  North  Atlantic 
and  Paratethys  regions  (see  Barnes,  1978:26).  The  family 
might  have  been  cosmopolitan  during  the  Miocene,  but  there 
are  as  yet  no  kentriodontids  recognized  from  the  South  At- 
lantic realm  or  from  the  eastern  South  Pacific.  It  would  be 
premature  to  speculate  on  the  place  of  origin  of  the  family. 


even  though  the  oldest  known  occurrence  is  in  the  South 
Pacific. 

Barnes  (1976:338)  noted  that  the  then  available  paleon- 
tologic  literature  suggested  a pattern  of  generic  cosmopoli- 
tanism among  larger  fossil  late  Tertiary  cetaceans  and  one 
of  generic  endemism  within  major  northern  ocean  basins 
among  the  smaller  fossil  odontocetes.  Among  the  nine  cur- 
rently recognized  genera  of  Kentriodontidae  (cf.  Barnes,  1978: 
26),  more  than  one-half  (i.e.  Liolithax  Kellogg,  1931,  Ken- 
triodon. Delphinodon  Leidy,  1 869,  Pithanodelphis  Abel,  1905, 
and  Lophocetus  Cope,  1868)  are  now  known  by  species  in 
both  the  North  Atlantic  and  the  North  Pacific  regions.  For 
the  Kentriodontidae,  it  now  appears  that  within  ocean  basins 
a pattern  of  endemism  at  the  species  level  rather  than  the 
generic  level  prevailed. 

PHYLOGENETIC  RELATIONSHIPS 

Kentriodon  is  the  most  primitive  genus  yet  assigned  to  the 
subfamily  Kentriodontidae.  For  example,  Kentriodon  lacks 
such  derived  characters  as  the  elevated  cranial  vertex  and 
enlarged  nasal  bones  of  Pithanodelphis.  and  the  short  ros- 
trum, bulbous  braincase  and  rounded  facial  margins  of  Del- 
phinodon dividum.  The  cranial  morphology  of  Kentriodon 
suggests  that  its  origin  might  have  been  among  more  prim- 
itive species  in  the  subfamily  Kampholophinae.  Kentriodon 
shares  with  the  kampholophine  species  Liolithax  pappus  and 
Kampholophos  serrulus  such  primitive  characters  as  a nar- 
row, elongate  rostrum,  high  tooth  count,  low  cranial  vertex, 
and  a concave  margin  of  the  facial  region  above  the  orbit. 
Kentriodon  is  more  derived,  however,  than  either  of  those 
species,  by  having  facial  surfaces  of  the  frontal  and  the  max- 
illa that  are  spread  more  over  the  dorsal  opening  of  the  tem- 
poral fossa.  The  extent  of  this  spreading,  however,  had  not 
progressed  to  the  stage  seen  in  the  derived  kentriodontine 
genera  Delphinodon,  Leptodelphis  Kirpichnikov,  1954,  Mi- 
crophocaena  Kudrin  and  Tatarinov,  1965,  Pithanodelphis 
and  Sarmatodelphis  Kirpichnikov,  1954. 

There  are  possibly  relationships  between  Kentriodontinae 
and  primitive  modem  Delphinidae,  such  as  species  in  the 
subfamily  Steninae  ( sensu  Mead,  1975),  and  this  has  been 
discussed  previously  by  True  (1912),  Kellogg  (1927),  and 
Barnes  (1978).  Similarities  exist  in  overall  cranial  propor- 
tions, shape  of  the  mandible,  and  size  and  numbers  of  teeth, 
but  kentriodontines  are  more  primitive  by  having  symmet- 
rical cranial  vertices,  less  extensive  air  sinuses  and  unfused 
cervical  vertebrae. 

The  two  essentially  synchronous  species,  Kentriodon  per- 
nix  and  K.  obscurus.  have  different  combinations  of  both 
primitive  and  derived  characters  (see  Table  1).  For  example, 
K.  pernix  is  more  derived  than  K.  obscurus  by  having  the 
premaxillary  foramina  located  more  posteriorly  (Fig.  1 3),  and 
the  posterior  part  of  the  palate  less  convex  on  either  side  of 
the  pterygoid  sinuses.  Kentriodon  obscurus.  on  the  other  hand, 
is  more  derived  by  having  shorter  postorbital  processes  of 
the  frontals,  smaller  teeth  (Fig.  14),  medial  premaxillary  sur- 


20  Contributions  in  Science,  Number  353 


Barnes  and  Mitchell:  Kentriodon  from  California 


face  in  front  of  the  nares  not  inclined,  a longer  anterior  ex- 
tension of  the  palatine  bones  on  the  palatal  surface  and  of 
the  pterygoid  air  sinuses  within  the  pterygoid  hamuli,  and  a 
larger  postorbital  lobe  of  the  pterygoid  air  sinus.  The  derived 
character  states  listed  in  Table  1 are  equally  spread  between 
the  two  species.  The  polarity  of  these  characters  was  deter- 
mined by  comparisons  with  species  in  the  more  primitive 
odontocete  families,  Squalodontidae  and  Agorophiidae. 

Kentriodon  pernix  and  K.  obscurus  might  have  evolved 
from  a common  ancestor,  and  the  discoveries  of  the  older 
fossils  of  kentriodontines  in  California  certainly  indicate  that 
such  Kentriodon- like  dolphins  lived  prior  to  Middle  Miocene 
time  in  the  North  Pacific  Ocean. 

CONCLUSIONS 

A fossil  delphinoid  genus,  Kentriodon  Kellogg,  1927,  that  is 
uncommon  in  northwest  Atlantic  Middle  Miocene  rocks, 
occurs  in  contemporaneous  rocks  on  the  eastern  margin  of 
the  North  Pacific  Ocean.  A few  fossils  from  the  North  Pacific 
margin  identified  as  this  genus  or  related  genera  of  small 
dolphins  have  been  briefly  cited  in  previous  literature,  and 
it  is  shown  here  that  the  genus  Grypolithax  Kellogg,  1931, 
originally  based  on  specimens  from  the  Middle  Miocene 
Sharktooth  Hill  Bonebed  in  California,  is  synonymous  with 
Kentriodon.  Based  on  isolated  fossil  periotics,  Kellogg  (1931) 
had  named  two  species  in  Grypolithax,  G.  obscura  Kellogg, 
1931,  and  G.  pavida  Kellogg,  1931.  These  are  the  only  species 
that  have  ever  been  assigned  to  Grypolithax  and  we  regard 
them  as  synonymous.  G.  obscura  has  page  priority,  is  the 
senior  synonym  of  G.  pavida,  and  is  the  type  species  of  the 
genus.  Because  Grypolithax  is  not  valid,  the  most  appropriate 
binomen  for  the  species  from  the  Sharktooth  Hill  Bonebed 
is  Kentriodon  obscurus  (Kellogg,  1931).  We  refer  periotics 
and  a partial  skull  to  this  species.  The  only  other  named 
species  of  Kentriodon  is  the  type  species,  K.  pernix  Kellogg, 
1927,  from  the  Calvert  Formation  in  Maryland. 

Among  possibly  as  many  as  1 4 species  of  odontocetes  found 
in  the  Sharktooth  Hill  Bonebed,  K.  obscurus  is  not  abundant 
and  is  one  of  the  rarest  species.  Only  one  of  the  rarer  ones, 
Platylithax  robusta  Kellogg,  1931,  has  been  named  previ- 
ously in  the  scientific  literature.  The  other  four  have  not  been 
named,  although  Barnes  (1976)  has  called  attention  to  their 
presence.  Only  by  increasing  the  total  sample  size  will  we  be 
able  to  eventually  recognize  the  rarer  species  in  the  Shark- 
tooth Hill  Local  Fauna  and  learn  their  anatomy. 

Middle  Miocene  fossils  of  Kentriodon  have  now  been  re- 
ported from  Maryland  and  California  in  the  U.S.A.,  and 
apparently  also  from  Japan.  The  earliest  reported  kentri- 
odontine  fossils  from  the  eastern  North  Pacific  region  are 
two  isolated  periotics  we  have  identified  as  cf.  Kentriodon 
sp.  from  the  early  Middle  Miocene  part  of  the  Round  Moun- 
tain Silt,  lower  stratigraphically  than  the  Sharktooth  Hill 
Bonebed.  Kentriodon  may  have  been  a cosmopolitan  genus 
in  the  Middle  Miocene,  but  there  is  yet  no  published  fossil 
evidence  from  the  southern  hemisphere  to  prove  this. 

Each  of  the  two  known  species  of  Kentriodon  has  a different 


suite  of  primitive  and  derived  characters,  and  the  two  are 
probably  derived  from  a common  ancestor. 

ACKNOWLEDGMENTS 

This  study  was  aided  by  the  granting  of  travel  funds  to  L.G. 
Barnes  by  the  Natural  History  Museum  of  Los  Angeles 
County  (LACM)  Foundation  and  the  Smithsonian  Institu- 
tion. Some  of  the  artwork  was  sponsored  by  the  University 
of  California  Museum  of  Paleontology  (UCMP).  Curation  of 
some  specimens  was  aided  by  National  Science  Foundation 
(NSF)  Grant  DEB  7202014  to  the  LACM  Foundation  for 
care  of  collections.  Early  fieldwork  by  E.D.  Mitchell  was 
supported  by  LACM.  Some  of  the  later  fieldwork  was  sup- 
ported by  NSF  Grant  EAR  7916508  to  the  LACM  Foun- 
dation to  facilitate  paleoecologic  investigations  of  the  Shark- 
tooth Hill  Bonebed. 

We  are  grateful  to  the  following  persons  for  collecting  var- 
ious specimens  used  in  this  study:  Mr.  and  Mrs.  E.L.  An- 
derson, Shelton  P.  Applegate,  Angela  Bennett,  Richard  C. 
Bishop,  Dennis  L.  Clayton,  the  late  John  E.  Fitch,  William 
Hawes,  Jr.,  Richard  W.  Huddleston,  Paul  Kirkland,  Douglas 
J.  Macdonald,  the  late  Charles  Morrice,  Michael  D.  Quarles, 
Camm  C.  Swift,  Terry  Story,  and  Howell  W.  Thomas. 

We  thank  Robert  L.  Brownell,  Jr.,  R.  Ewan  Fordyce,  Sam- 
uel A.  McLeod,  James  G.  Mead,  David  P.  Whistler,  Frank 
C.  Whitmore,  Jr.,  and  anonymous  reviewers  for  constructive 
criticism  of  the  manuscript. 

The  following  persons  allowed  us  to  study  specimens  in 
their  care:  Clayton  E.  Ray,  Frank  C.  Whitmore,  Jr.,  and 
Robert  Purdy  (USNM),  Victor  A.  Zullo  and  Peter  Rodda 
(CAS),  and  J.  Howard  Hutchison  (UCMP). 

We  were  kindly  granted  access  to  collecting  sites  in  Kern 
County  by  Mr.  and  Mrs.  Clint  Smoot,  Mr.  and  Mrs.  Steven 
Smoot,  Vincent  Bertolucci,  the  late  Bernard  Terabino,  and 
E.H.  Shuler  and  other  employees  of  Getty  Oil  Company. 

David  Cook,  formerly  of  UCMP,  prepared  some  of  the 
line  drawings  (Fig.  1 1).  The  photographs  (Figs.  2,  4,  6,  8,  9) 
were  prepared  by  Lawrence  Reynolds,  formerly  of  LACM. 
John  De  Leon  of  LACM  did  some  of  the  photographic  copy- 
ing of  the  artwork.  Other  illustrations  were  prepared  by  L.G. 
Barnes. 

LITERATURE  CITED 

Abel,  O.  1905.  Les  odontocetes  du  Bolderien  (Miocene 
superieur)  d’Anvers.  Memoires  du  Musee  Royal  d’His- 
toire  Naturelle  de  Belgique  3(2):  1-1  55. 

Addicott,  W.O.  1972.  Provincial  middle  and  late  Tertiary 
molluscan  stages.  Temblor  Range,  California.  Pages  1- 
26  in  E.H.  Stinemeyer,  ed..  Proceedings  of  the  Pacific 
Coast  Miocene  Biostratigraphic  Symposium,  Society  of 
Economic  Paleontologists  and  Mineralogists.  364  pp. 
Barnes,  L.G.  1976.  Outline  of  eastern  North  Pacific  fossil 
cetacean  assemblages.  Systematic  Zoology  25(4):32 1 — 
343. 


Contributions  in  Science,  Number  353 


Barnes  and  Mitchell:  Kentriodon  from  California  21 


. 1978.  A review  of  Lophocetus  and  Liolithax  and 

their  relationships  to  the  delphinoid  family  Kentriodon- 
tidae  (Cetacea:  Odontoceti).  Natural  History  Museum 
of  Los  Angeles  County  Science  Bulletin  28:1-35. 

Berggren,  W.,  and  J.A.  Van  Couvering.  1974.  The  late  Neo- 
gene. Biostratigraphy,  geochronology  and  paleoclima- 
tology  of  the  last  15  million  years  in  marine  and  con- 
tinental sequences.  Palaeogeography,  Palaeoclimatology 
and  Palaeoecology  16(1  /2):i— xi,  1-216. 

Boenninghaus,  G.  1904.  Das  Ohr  des  Zahnwales,  zugleich 
ein  Beitrag  zur  Theorie  der  Schalleitung.  Eine  biolo- 
gische  Studie.  Zoologische  Jahrbiicher.  Abteilung  fur 
Anatomie  und  Ontogenie  der  Tiere  1 9: 1 89-360,  pis.  1 2- 
13. 

Clark,  W.B.,  G.B.  Shattuck,  and  W.H.  Dali.  1904.  The 
Miocene  Deposits  of  Maryland.  Pages  xxi-xxxii  in  W.B. 
Clark,  ed.,  Maryland  Geological  Survey— Miocene,  Bal- 
timore, The  Johns  Hopkins  Press. 

Cope,  E.D.  1868a  (for  1867).  (Descriptions  of  Eschrichtius 
cephalus,  Rhabdosteus  latiradix,  Squalodon  atlanticus 
and  S.  mento.)  Proceedings  of  the  Academy  of  Natural 
Sciences,  Philadelphia  19:131-132. 

. 1868b  (for  1867).  An  addition  to  the  vertebrate 

fauna  of  the  Miocene  period,  with  a synopsis  of  the 
extinct  Cetacea  of  the  United  States.  Proceedings  of  the 
Academy  of  Natural  Sciences,  Philadelphia  19: 1 38-156. 

Denker,  A.  1902.  Zur  Anatomie  des  Gehororgans  der  Ce- 
tacea. Anatomische  Hefte  19(2):423-448,  pis.  14-15. 

Evemden,  J.F.,  D.E.  Savage,  G.H.  Curtis,  and  G.T.  James. 
1964.  Potassium-argon  dates  and  the  Cenozoic  mam- 
malian chronology  of  North  America.  American  Journal 
of  Science  262:145-198. 

Fordyce,  R.E.  1980.  Whale  evolution  and  Oligocene  south- 
ern ocean  environments.  Palaeogeography,  Palaeocli- 
matology and  Palaeoecology  31:319-336. 

. 1981.  Systematics  of  the  odontocete  whale  Ago- 

rophius  pygmaeus  and  the  family  Agorophiidae  (Mam- 
malia: Cetacea).  Journal  of  Paleontology  5 5(5):  1 028— 
1045. 

Fraser,  F.C.,  and  P.E.  Purves.  1960.  Hearing  in  cetaceans. 
Evolution  of  the  accessory  air  sacs  and  the  structure  and 
function  of  the  outer  and  middle  ear  in  Recent  cetaceans. 
Bulletin  of  the  British  Museum  (Natural  History),  Zo- 
ology 7(1):  1-140,  frontispiece,  pis.  1-53. 

Gazin,  C.L.,  and  R.L.  Collins.  1950.  Remains  of  land  mam- 
mals from  the  Miocene  of  the  Chesapeake  Bay  region. 
Smithsonian  Miscellaneous  Collections  1 16(2):  1-21. 

Hay,  O.P.  1902.  Bibliography  and  catalogue  of  the  fossil 
Vertebrata  of  North  America.  Bulletin  of  the  United 
States  Geological  Survey  179:1-868. 

Kasuya,  T.  1973.  Systematic  consideration  of  Recent 
toothed  whales  based  on  the  morphology  of  tympano- 
periotic  bone.  Scientific  Reports  of  the  Whales  Research 
Institute  25:1-103,  pis.  1-28. 

Kellogg,  A.R.  1925.  The  relationships  of  the  Tertiary  ce- 
taceans of  Jugo-Slavia  to  those  of  eastern  North  Amer- 
ica. Spomenice  U Pocast  Prof.  Dr.  Gorjanovic-Kram- 
bergera,  Zagreb,  pp.  1-8. 


. 1927a.  Kentriodon  pernix,  a Miocene  porpoise  from 

Maryland.  Proceedings  of  the  United  States  National 
Museum  69(  1 9):  1 —55,  pis.  1-14. 

. 1927b.  Study  of  the  skull  of  a fossil  sperm-whale 

from  the  Temblor  Miocene  of  Southern  California.  Con- 
tributions to  Paleontology,  Carnegie  Institution  of 
Washington  Publications  346:1-24,  pis.  1-9. 

. 1928.  The  history  of  whales— their  adaptation  to 

life  in  the  water.  Quarterly  Review  of  Biology  3(1,  2): 
29-76,  174-208. 

. 1931.  Pelagic  mammals  from  the  Temblor  For- 
mation of  the  Kern  River  region,  California.  Proceedings 
of  the  California  Academy  of  Sciences,  Ser.  4,  19(12): 
217-397. 

. 1955.  Three  Miocene  porpoises  from  the  Calvert 

Cliffs,  Maryland.  Proceedings  of  the  United  States  Na- 
tional Museum  105:101-154,  pis.  1-21. 

Kellogg,  A.R.,  and  F.C.  Whitmore,  Jr.  1957.  Mammals. 
Pages  1021-1024  in  H.S.  Ladd,  ed..  Treatise  on  marine 
ecology  and  paleoecology.  Volume  2,  Paleoecology. 
Geological  Society  of  America  Memoir  67:i-x,  1-1077. 

Kirpichnikov,  A. A.  1954.  Dva  novykh  roda  del’finov  iz 
sarmata  SSSR.  (Two  new  genera  of  dolphins  from  Sar- 
matian  of  U.S.S.R.)  Trudy  Paleontologicheskogo  Insti- 
tuta,  Akademia  Nauk  S.S.S.R.  47:181-190,  pis.  1-3. 

Kudrin,  L.N.,  and  K. A.  Tatarinov.  1965.  O miotzenovykh 
del’finakh  zapadnoi  Ukrainy.  (On  the  Miocene  delphin- 
ids  from  the  western  Ukraine.)  Paleontologicheskii 
Zhumal  1 965(4):68-74. 

Lawrence,  B.,  and  W.E.  Schevill.  1956.  The  functional 
anatomy  of  the  delphinid  nose.  Bulletin  of  the  Museum 
of  Comparative  Zoology,  Harvard  College  1 1 4(4):  1 03— 
151. 

Leidy,  J.  1869.  The  extinct  mammalian  fauna  of  Dakota 
and  Nebraska,  including  an  account  of  some  allied  forms 
from  other  localities,  together  with  a synopsis  of  the 
mammalian  remains  of  North  America.  Journal  of  the 
Academy  of  Natural  Sciences  of  Philadelphia,  new  series 
7:1-472. 

Mead,  J.G.  1975.  Anatomy  of  the  external  nasal  passages 
and  facial  complex  in  the  Delphinidae  (Mammalia:  Ce- 
tacea). Smithsonian  Contributions  in  Zoology  207:i~iv, 
1-72. 

Miller,  G.S.,  Jr.  1923.  The  telescoping  of  the  cetacean  skull. 
Smithsonian  Miscellaneous  Collections  76(5):  1-71. 

Mitchell,  E.  1965.  History  of  research  at  Sharktooth  Hill, 
Kern  County,  California.  Kern  County  Historical  So- 
ciety (Bakersfield,  California),  Special  Publication,  vi  + 
45  pp. 

. 1966.  The  Miocene  pinniped  Allodesmus.  Univer- 
sity of  California  Publications  in  Geological  Sciences  6 1 : 
1-46,  pis.  1-29. 

, and  R.H.  Tedford.  1973.  The  Enaliarctinae,  a new 

group  of  extinct  aquatic  Carnivora  and  a consideration 
of  the  origin  of  the  Otariidae.  Bulletin  of  the  American 
Museum  of  Natural  History  151(3):201  —284. 

Okazaki,  Y.  1976.  Miocene  long-snouted  porpoises  from 


22  Contributions  in  Science,  Number  353 


Barnes  and  Mitchell:  Kentriodon  from  California 


the  Mizunami  Group,  central  Japan.  Bulletin  of  the  Mi- 
zunami  Fossil  Museum  3:25-40,  pis.  9-1 1. 

Ray,  C.E.  1976.  Geography  of  phocid  evolution.  Systematic 
Zoology  25(4):39 1-406. 

Rensberger,  J.M.  1 969.  A new  iniid  cetacean  from  the  Mio- 
cene of  California.  University  of  California  Publications 
in  Geological  Sciences  82:1-34,  pis.  1-4. 

Repenning,  C.A.,  and  R.H.  Tedford.  1977.  Otarioid  seals 
of  the  Neogene.  United  States  Geological  Survey  Profes- 
sional Paper  992:i-vi,  1-93,  pis.  1-24. 

Savage,  D.E.,  and  L.G.  Barnes.  1972.  Miocene  vertebrate 
geochronology  of  the  west  coast  of  North  America.  Pages 
124-145  in  E.H.  Stinemeyer,  ed.,  Proceedings  of  the 
Pacific  Coast  Miocene  Biostratigraphic  Symposium,  So- 
ciety of  Economic  Paleontologists  and  Mineralogists.  364 
PP- 

Shikama,  T.,  Y.  Hasegawa,  and  H.  Otsuka.  1973.  Geolog- 
ical range  of  mammals  in  the  Japanese  Neogene.  Mem- 
oirs of  the  Geological  Society  of  Japan  8:137-141  (in 
Japanese,  English  abstract). 


Simpson,  G.G.,  A.  Roe,  and  R.C.  Lewontin.  1960.  Quan- 
titative Zoology.  Revised  Edition.  Harcourt,  Brace  and 
World,  Inc.,  New  York,  pp.  i-vii,  1-440. 

Slijper,  E.J.  1936.  Die  Cetaceen.  Vergleichend-anatomisch 
und  systematisch.  Capita  Zoologica  6:i-xv,  1-590. 

. 1958.  Walvissen.  D.B.  Centen’s  Uitgeversmaat- 

schappij,  Amsterdam.  524  pp. 

True,  F.W.  1912.  Description  of  a new  fossil  porpoise  of 
the  genus  Delphinodon  from  the  Miocene  Formation  of 
Maryland.  Journal  of  the  Academy  of  Natural  Sciences 
of  Philadelphia,  Ser.  2,  15:165-194,  pis.  17-26. 

Weaver,  C.E.  et  al.  1944.  Correlation  of  the  marine  Ce- 
nozoic  formations  of  western  North  America.  Geolog- 
ical Society  of  America  Bulletin  55:569-598;  pi.  1. 

Wood,  H.E.  et  al.  1941.  Nomenclature  and  correlation  of 
the  North  American  continental  Tertiary.  Geological  So- 
ciety of  America  Bulletin  52:1-48,  pi.  1. 

Accepted  for  publication  4 April  1984. 


Contributions  in  Science,  Number  353 


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SYSTEMATICS  OF  FISSURELLA  IN  THE  PERUVIAN 
AND  MAGELLANIC  FAUNAL  PROVINCES 
(GASTROPODA:  PROSOBRANCHIA) 

James  H.  McLean 


Contributions  in  Science,  Number  354 
Natural  History  Museum  of  Los  Angeles  County 
29  October  1984 


ISSN  0459-8113 

Natural  History  Museum  of  Los  Angeles  County 
900  Exposition  Boulevard 
Los  Angeles,  California  90007 


CONTENTS 


ABSTRACT  1 

RESUMEN 1 

INTRODUCTION  2 

MATERIALS  AND  METHODS 3 

Fieldwork 3 

Other  Collections  Examined  3 

Preparation  of  Specimens 3 

Conventions  3 

Abbreviations 3 

HISTORICAL  REVIEW  4 

STRUCTURE 5 

Internal  Anatomy 5 

External  Anatomy  6 

Radula  6 

Shell  Morphology 8 

Shell  Structure 8 

BIOLOGY  AND  ECOLOGY 8 

Habitat 8 

Feeding 8 

Reproduction  and  Growth 9 

Epibiotic  Associations 9 

Epibiotic  Scurria  parasitica 9 

Shell  Borers 9 

Parasites 1 1 

Seastar  Predators 11 

Vertebrate  Predators 1 1 

Human  Predation  and  Economic  Importance 11 

SYSTEMATIC  CHARACTERS 12 

Size  and  Shape 12 

Sculpture  1 2 

Color  Pattern 1 2 

Interior  Margin 1 2 

Foramen 1 3 

Juvenile  Shell  13 

External  Anatomy  13 

Radula  13 

NAMES,  ALLOCATIONS,  AND  KEY  CHARACTERS 13 

Names  and  Allocations 13 

Chart  of  Key  Characters 14 

DISTRIBUTION  AND  ZOOGEOGRAPHY 15 

FOSSIL  RECORD 15 

FORMAT  FOR  SPECIES  ACCOUNTS 16 

SYSTEMATICS 17 

Family  Fissurellidae  Fleming,  1822  17 

Subfamily  Fissurellinae  Fleming,  1822  18 

Genus  Fissurella  Bruguiere,  1789 18 

Subgenus  Fissurella  Bruguiere.  1789  19 

Group  of  F.  peruviana 19 

Fissurella  peruviana  Lamarck,  1822 21 

Group  of  F.  maxima 24 

Fissurella  maxima  Sowerby,  1835  25 

Fissurella  latimarginata  Sowerby,  1835  28 

Fissurella  cumingi  Reeve,  1849 31 

Fissurella  costata  Lesson,  1831 34 

Fissurella  picta  (Gmelin,  1791) 37 


Fissurella  radiosa  Lesson,  1831 43 

Fissurella  oriens  Sowerby,  1835  49 

Fissurella  nigra  Lesson,  1831  52 

Group  of  F.  limbata  55 

Fissurella  limbata  Sowerby,  1835 55 

Fissurella  crassa  Lamarck,  1822  58 

Fissurella  bridgesii  Reeve,  1 849  60 

Fissurella  pulchra  Sowerby,  1835  63 

LACM  LOCALITIES  FOR  FIGURED  SPECIMENS 65 

ACKNOWLEDGMENTS  66 

LITERATURE  CITED 67 


SYSTEMATICS  OF  FISSURELLA  IN  THE  PERUVIAN  AND 
MAGELLANIC  FAUNAL  PROVINCES 
(GASTROPODA:  PROSOBRANCHIA) 

James  H.  McLean1 


ABSTRACT.  Fifty-eight  names  have  been  proposed  for  the  large 
and  abundant  species  of  Fissurella  in  the  Peruvian  and  Magellanic 
faunal  provinces.  Fieldwork  in  Peru  and  Chile  and  to  a lesser  extent 
in  Argentina,  has  produced  large  collections.  Following  study  of  these 
collections,  as  well  as  most  of  the  type  specimens,  I reduce  the 
number  of  species  to  1 3,  three  of  which  have  geographic  subspecies. 

Peruvian-Magellanic  species  of  Fissurella  Bruguiere,  1789,  are 
members  of  the  nominate  subgenus,  in  which  the  shell  has  an  inner 
layer  of  crossed  lamellar  aragonite  and  a thick  outer  layer  of  prismatic 
calcite.  In  contrast,  most  tropical  species  of  Fissurella,  which  are  in 
the  subgenus  Cremides  H.  and  A.  Adams,  1854,  have  a shell  com- 
posed entirely  of  aragonite.  The  outer  layer  of  calcite  is  evidently  an 
adaptation  to  cold  water  in  the  Peruvian-Magellanic  species. 

Three  species  groups  in  Fissurella  ( sensu  stricto)  are  recognized. 
The  group  of  Fissurella  peruviana  Lamarck,  1822,  is  smaller-shelled 
and  has  a thinner  calcitic  layer  than  species  in  the  other  groups;  this 
group  includes  the  type  species  F.  nimbosa  Linnaeus,  1758,  in  the 
southern  Caribbean  (the  only  tropical  member  of  the  subgenus)  and 
F.  volcano  Reeve,  1849,  in  California  and  Baja  California. 

The  group  of  F.  maxima  Sowerby,  1 833,  is  characterized  by  strong 
primary  and  secondary  ribs  (at  least  in  juvenile  stages)  and  also 
includes  F.  latimarginata  Sowerby,  1835,  F.  cumingi  Reeve,  1849, 
and  F.  costata  Lesson,  1831,  in  the  Peruvian  Province,  and  F.  picla 
(Gmelin,  1791),  F.  radiosa  Lesson,  1831,  F.  oriens  Sowerby,  1835, 
and  F.  nigra  Lesson,  1831,  in  the  Magellanic  Province.  The  group 
of  F.  limbata  Sowerby,  1835,  is  characterized  by  broad  primary  nbs 
and  lack  of  secondary  ribs;  it  includes  F.  crassa  Lamarck,  1 822,  F. 
bridgesii  Reeve,  1849,  and  F.  pulchra  Sowerby,  1835,  all  in  the 
Peruvian  Province. 

Geographic  subspecies  are  here  recognized  for  three  species  broad- 
ly distributed  in  the  Magellanic  Province:  F.  picta  picta  (Gmelin, 
1791)  in  southern  Chile,  and  F.  picta  lata  Sowerby,  1835,  in  central 
Chile;  F.  radiosa  radiosa  Lesson,  1831,  in  southern  Chile,  F.  radiosa 
tixierae  Metivier,  1969,  in  the  vicinity  of  the  Gulf  of  San  Matias  in 
Argentina;  F.  oriens  oriens  Sowerby,  1835,  in  southern  Chile,  and 
F.  oriens  fulvescens  Sowerby,  1835,  in  central  Chile. 

Three  species,  F.  cumingi,  F.  bridgesii,  and  F.  pulchra,  have  been 
poorly  understood  by  previous  authors  and  are  newly  defined  here. 

Distributions  of  the  Peruvian  and  Magellanic  species  overlap  in 
south-central  Chile,  where  1 2 of  the  1 3 species  occur. 

This  account  includes  observations  on  shell  epibionts  and  borers, 
and  reviews  the  sparse  literature  on  the  biology  of  these  species. 

Contributions  in  Science,  Number  354,  pp.  1-70 
Natural  History  Museum  of  Los  Angeles  County,  1984 


RESUMEN.  Hasta  ahora  habian  sido  propuestos  58  nombres  para 
las  grandes  y abundantes  especies  de  Fisssurella  de  las  provincias 
biogeograficas  Peruana  y Magallanica.  Trabajos  de  terrene  en  Peru 
y Chile,  y en  menor  intensidad  en  Argentina,  han  proporcionado 
importantes  colecciones.  Realizado  el  estudio  de  estas  colecciones  y 
de  muchos  de  los  ejemplares  tipo,  se  reduce  a 13  el  numero  de 
especies,  tres  de  las  cuales  poseen  subespecies  geograficas. 

Las  especies  de  Fissurella  de  las  provincias  Peruana  y Magallanica 
son  integrantes  del  subgenero  Fissurella  Bruguiere,  1789  {sensu  stric- 
to), en  las  cuales  la  concha  tiene  una  capa  interna  compuesta  de 
aragonita  laminar  cruzada  y una  capa  mas  externa  de  calcita  pris- 
matica.  En  cambio,  muchas  especies  de  Fissurella  que  pertenecen 
al  subgenero  Cremides  H.  y A.  Adams,  1854,  tienen  la  concha  com- 
puesta integramente  de  aragonita.  La  capa  mas  externa  de  calcita  es 
considerada  como  una  adaptacion  de  las  especies  de  las  provincias 
Peruana  y Magallanica  a aguas  frias. 

Se  reconocen  tres  grupos  de  especies.  Un  primer  grupo  de  Fis- 
surella peruviana  Lamarck,  1822,  de  concha  mas  pequena  y cuya 
capa  de  calcita  es  mas  delgada  que  en  las  otras  especies  de  los  demas 
grupos.  Este  grupo  comprende  la  especies  tipo  F.  nimbosa  Linnaeus, 
1758,  del  sur  del  Caribe  (unico  miembro  tropical  del  subgenero)  y 
F.  volcano  Reeve,  1849,  de  California  y Baja  California. 

Un  segundo  grupo  de  F.  maxima  Sowerby,  1833,  caracterizado 
por  la  presencia  de  gruesas  costillas  primarias  y secundarias  (al  me- 
nos  en  los  estados  juveniles).  Comprende  tambien  a F.  latimargi- 
nata Sowerby,  1835,  F.  cumingi  Reeve,  1849,  y F.  costata  Lesson, 
1 83 1 , de  la  provincia  Peruana  y a F.  picta  (Gmelin,  1 79 1 ),  F.  radiosa 
Lesson,  1831,  F.  oriens  Sowerby,  1835,  y F.  nigra  Lesson,  1831,  de 
la  provincia  Magallanica. 

El  tercer  grupo  de  F.  limbata  Sowerby,  1835,  esta  caracterizado 
por  la  presencia  de  costillas  primanas  anchas  y ausencia  de  costillas 
secundarias.  Comprende  tambien  a F.  crassa  Lamarck,  1822,  F. 
bridgesii  Reeve,  1849,  y F.  pulchra  Sowerby,  1835,  todas  de  la  pro- 
vincia Peruana. 

Se  reconocen  las  siguientes  subespecies  geograficas  para  tres  es- 
pecies ampliamente  distribuidas  en  la  provincia  Magallanica:  F. 
picta  picta  (Gmelin,  1791)  del  sur  de  Chile  y F.  picta  lata  Sowerby, 
1835,  de  la  zona  central  de  Chile;  F.  radiosa  radiosa  Lesson,  1831, 


1.  Malacology  Section,  Natural  History  Museum  of  Los  Angeles 
County,  900  Exposition  Blvd.,  Los  Angeles,  California  90007. 


ISSN  0459-81  13 


Figure  1.  A Fissurella  “shell  pile,”  consisting  of  large  specimens  of  F.  latimarginata,  F.  cumingi.  and  F.  maxima  at  Los  Molles,  Aconcagua 
Province,  Chile,  October  15,  1975.  The  specimens  had  presumably  been  taken  by  shallow  diving  in  the  vicinity  and  the  shells  discarded.  All 
shells  were  covered  with  the  algal  mat,  which  completely  obscures  the  color  pattern. 


del  sur  de  Chile  y F.  radiosa  tixierae  Metivier,  1 969,  de  las  cercanias 
del  golfo  de  San  Matias  en  Argentina;  F.  oriens  oriens  Sowerby, 
1835,  del  sur  de  Chile  y F.  oriens  fulvescens  Sowerby,  1 835,  de  Chile 
central. 

Las  especies  F.  cumingi,  F.  bridgesii  y F.  pulchra,  que  han  sido 
escasamente  tratadas  por  autores  anteriores,  son  definidas  nueva- 
mente. 

Se  observa  una  sobreposicion  en  la  distribucion  de  las  especies  de 
las  provincias  Peruana  y Magallanica  en  la  zona  central-sur  de  Chile, 
donde  1 2 de  las  13  especies  se  encuentran  presentes. 

En  el  presente  trabajo  se  incluye,  ademas,  observaciones  sobre  los 
epibiontes  y organismos  perforadores  de  las  conchas  y se  revisa  la 
esparcida  literatura  que  trata  sobre  la  biologia  de  algunas  de  estas 
especies. 

INTRODUCTION 

The  Fissurella  species  of  the  cool  waters  of  Peru,  Chile,  and 
southern  Argentina  are  large  and  abundant,  comprising  a 
major  element  of  the  mollusk  fauna  of  the  west  coast  of  South 
America.  They  are  extensively  used  for  food  and  are  known 
locally  as  “lapas”  (Fig.  1).  The  importance  of  the  fishery  is 
second  only  to  that  of  Concholepas,  the  “loco,”  the  large 
limpetlike  thaidid  gastropod  of  the  region.  Despite  this  im- 
portance, the  taxonomy  of  the  South  American  species  of 
Fissurella  has  been  poorly  understood. 

It  has  been  recognized  that  a large  number  of  highly  vari- 
able, sympatric  species  occur  in  the  region.  Some  58  names 


for  Recent  species  have  been  introduced  in  the  literature. 
Widely  varying  estimates  of  the  number  of  actual  species 
have  been  given:  Pilsbry  (1890)  recognized  about  20  species; 
Ziegenhom  and  Thiem  (1925)  treated  1 1 species  and  three 
“varieties”;  Riveros-Zuniga  (1951)  recognized  26  species  and 
three  “varieties”;  Dell  (1971)  listed  1 1 possible  species;  and 
finally  Ramirez-Boehme  (1974)  gave  a key  to  30  species  and 
two  “varieties.” 

The  collection  of  mollusks  from  Iquique,  Chile,  reported 
upon  by  Marincovich  (1973),  and  deposited  in  the  Natural 
History  Museum  of  Los  Angeles  County,  included  five  species 
of  Fissurella.  Although  this  material  introduced  me  to  the 
subject,  fieldwork  of  my  own  in  Peru  in  1972  and  1974,  and 
in  Chile  for  two  months  in  1975,  and  southern  Argentina  in 
1978,  enabled  me  to  collect  and  observe  the  Fissurella  species 
from  many  different  localities.  I have  therefore  been  able  to 
observe  these  mollusks  throughout  their  entire  geographic 
range  from  north-central  Peru  to  southern  Chile  and  Argen- 
tina. 

The  Magellanic  Province  of  southern  Chile  and  southern 
Argentina  is  also  the  center  of  distribution  of  another  fissu- 
rellid  group  comprising  the  species  Fissurellidea  megatrema 
Orbigny,  1841,  F.  patagonica  (Strebel,  1907),  Pupillaea  an- 
nulus (Odhner,  1932),  and  the  shell-less  Buchanania  onchi- 
dioides  Lesson,  1830.  A report  on  these  species  has  been 
published  (McLean,  1984b). 


2 Contributions  in  Science,  Number  354 


McLean;  Peruvian  and  Magellanic  Fissurella 


In  this  work  I offer  a revised  classification  of  the  South 
American  species  of  Fissurella,  based  on  my  field  observa- 
tions, study  of  the  large  collection  now  housed  at  the  Los 
Angeles  County  Museum  of  Natural  History,  and  study  of 
type  material  borrowed  from  other  museums.  The  classifi- 
cation cannot  be  exhaustive  and  does  not  offer  a cladistic 
hypothesis  of  relationships.  In  the  absence  of  anatomical, 
biochemical  (electrophoretic),  and  other  characters,  that  is 
beyond  the  scope  of  the  present  work.  It  is  hoped  that  this 
paper  will  provide  a basis  for  future  work  on  the  systematics 
and  ecology  of  these  species. 

MATERIALS  AND  METHODS 

Fieldwork 

The  collection  upon  which  this  report  is  chiefly  based  is  now 
in  the  Los  Angeles  County  Museum  of  Natural  History.  Some 
material  from  miscellaneous  sources  is  represented  in  the 
collection,  but  the  bulk  of  it  resulted  from  nine  major  ex- 
peditions as  follows: 

1.  Peru:  Isla  San  Lorenzo,  Isla  Chinchas,  and  Bahia  Inde- 
pendencia.  Allan  Hancock  Expeditions,  January,  1935, 
and  February,  1938,  intertidal  and  dredging  stations. 

2.  Chile:  Iquique.  Louie  Marincovich,  June-September, 
1964,  June-July,  1970,  intertidal  stations. 

3.  Argentina:  Isla  de  los  Estados  (E  of  Tierra  del  Fuego).  R/V 
HERO,  April,  1971,  and  October,  1971,  intertidal  and 
dredging  stations;  collections  received  from  the  Smith- 
sonian Oceanographic  Sorting  Center. 

4.  Peru:  Pucasana,  Laguna  Grande,  Isla  Chincha  Norte,  Par- 
acas,  Asia.  James  H.  McLean,  April,  1972,  intertidal  and 
diving  stations. 

5.  Chile:  south  of  Isla  de  Chiloe.  Paul  Dayton,  on  R/V  HERO, 
October-November,  1972,  intertidal  and  diving  stations. 

6.  Chile  and  Argentina:  Strait  of  Magellan  and  Isla  de  los 
Estados.  Paul  Dayton,  on  R/V  HERO,  May,  1973,  inter- 
tidal and  diving  stations. 

7.  Peru:  Isla  Guanape,  Ancon,  and  Isla  San  Lorenzo.  James 
H.  McLean,  January,  1974,  intertidal  and  diving  stations. 

8.  Chile:  Iquique,  Antofagasta,  Coquimbo,  Los  Molles, 
Montemar,  Cartagena,  Concepcion,  Mehuin,  Pargua, 
Guabun,  Pumalin,  Islota  Nihuel,  Isla  Laitec,  Puerto 
Hambre,  Punta  Arenas.  James  H.  McLean,  October-No- 
vember, 1975,  intertidal  and  diving  stations. 

9.  Argentina:  Golfo  Nuevo  and  Golfo  San  Jose.  James  H. 
McLean,  on  R/V  HERO,  July  1978,  intertidal  and  dredg- 
ing stations. 

Other  Collections  Examined 

Upon  returning  from  Chile  in  1975,  I compared  the  field- 
collected  specimens  with  as  many  of  the  types  of  species 
described  by  nineteenth-century  authors  as  could  be  located 
for  me  in  the  British  Museum  (Natural  History)  and  the  Paris 
Museum.  I have  also  studied  the  collections  of  Fissurella  in 
the  U.S.  National  Museum  of  Natural  History,  Washington, 
D.C.,  the  Academy  of  Natural  Sciences,  Philadephia,  and  the 
American  Museum  of  Natural  History,  New  York.  Other 


specimens  were  received  on  loan  from  the  Museum  of  Com- 
parative Zoology,  Harvard,  and  the  National  Museum  of 
New  Zealand,  Wellington.  After  conducting  my  fieldwork  in 
Argentina  in  1978,  I examined  the  Fissurella  material  in  the 
Museo  Argentino  de  Ciencias  Naturales,  Buenos  Aires,  and 
the  Museo  Nacional  de  Historia  Natural,  Santiago.  Records 
from  these  collections  enabled  further  refinements  in  species 
distributions. 

Preparation  of  Specimens 

Although  most  of  the  field-collected  specimens  were  either 
kept  dry  or  the  entire  specimen  preserved  in  alcohol,  without 
cleaning  the  shell,  the  photographed  specimens  had  to  be 
cleaned  of  encrusting  organisms.  Shells  were  placed  in  full 
strength  laundry  bleach,  which  softens  the  algal  mat  and 
loosens  other  encrusting  organisms  so  that  the  shell  can  be 
scraped  clean  with  a knife  or  wire  brush.  Color  was  restored 
with  a light  application  of  mineral  oil. 

Shells  of  each  species  were  embedded  in  plaster  and  cut 
with  a diamond  rock  saw  for  the  examination  and  photog- 
raphy of  the  shell  layers.  Scanning  electron  microscopy  (SEM) 
was  used  for  the  examination  of  shell  structure  in  a fragment 
of  a small  specimen  of  F.  latimarginata.  Radulae  of  large 
specimens  of  each  species  were  air-dried  for  macrophotog- 
raphy; radulae  of  small  specimens  were  prepared  for  both 
light  microscopy  and  SEM. 

Conventions 

Figured  specimens  for  each  species  are  arranged  by  localities 
from  north  to  south,  including  type  specimens  of  nominate 
taxa  and  synonyms.  All  shell  specimens  are  illustrated  with 
the  anterior  at  the  top;  lateral  views  are  those  of  the  left  side 
of  the  shell.  Measurements  for  the  figured  specimens  are 
given  in  the  captions,  not  repeated  in  the  text.  Measurements 
are  given  in  this  order:  length,  width,  and  height.  Unless 
otherwise  indicated,  the  figured  specimens  were  collected  in 
the  intertidal  zone.  Latitude  and  longitude  for  the  figured 
specimens  from  LACM  stations  are  given  in  a locality  list 
following  the  systematic  section. 

Abbreviations 

Abreviations  of  institutions  mentioned  in  the  text  are  as 
follows: 

AHF  Allan  Hancock  Foundation  Collection  (at  LACM) 
AMNH  American  Museum  of  Natural  History,  New  York 

ANSP  Academy  of  Natural  Sciences,  Philadelphia 
BMNH  British  Museum  (Natural  History),  London 
LACM  Los  Angeles  County  Museum  of  Natural  History, 
Los  Angeles 

MACN  Museo  Argentino  de  Ciencias  Naturales,  Buenos 
Aires 

MCZ  Museum  of  Comparative  Zoology,  Harvard  Uni- 

versity, Cambridge 

MNHN  Museo  Nacional  de  Historia  Natural,  Santiago 
MNHNP  Museum  National  d’Histoire  Naturelle,  Paris 
NMNZ  National  Museum  of  New  Zealand,  Wellington 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  3 


USNM  United  States  National  Museum  of  Natural  His- 
tory, Washington,  D.C. 

HISTORICAL  REVIEW 

The  Strait  of  Magellan  was  probably  the  origin  of  the  earliest 
collected  shells  of  Fissurella  to  reach  Europe.  The  first  species 
to  be  known  was  the  common  Magellanic  F.  picta,  which 
was  well  described  and  figured  in  the  non-binomial  work  of 
Davila  (1767),  who  called  it  “Un  Lepas  rare  de  Magellan” 
(Pilsbry,  1890:114).  The  name  Fissurella  picta  dates  from 
J . F.  Gmelin  (1791),  whose  knowledge  of  it  came  in  part  from 
Davila. 

J.B.P.A.  de  Lamarck  (1822)  validated  F.  crassa  and  F. 
peruviana  in  his  “Histoire  naturelle  des  animaux  sans  ver- 
tebres.”  Lamarck’s  types  are  preserved  at  the  Geneva  Mu- 
seum (Mermod,  1950). 

G.B.  Sowerby  (1825)  introduced  a synonym  for  F.  crassa, 
but  was  later  to  recognize  Lamarck’s  unfigured  species. 

G.P.  Deshayes  (1830)  named  two  Chilean  species  in  the 
“Encyclopedic  Methodique,”  but  one  of  them  was  a synonym 
of  F.  peruviana  and  the  other,  F.  rudis,  although  prior  to  F. 
cost  at  a Lesson,  1831,  is  preoccupied.  Types  are  extant  at  the 
Paris  Museum. 

R.P.  Lesson  (1831),  in  his  report  on  collections  from  the 
South  American  voyage  of  the  “Coquille,”  described  four 
species  of  Fissurella  without  illustrations,  all  from  the  south- 
ern and  southernmost  regions  of  Chile.  The  names  for  three 
of  his  species  are  now  in  use:  F.  nigra,  F.  radiosa  and  F. 
costata.  One  other,  F.  obovalis,  remains  a nomen  dubium. 
Lesson’s  work  has  frequently  been  cited  as  published  in  1830, 
but  the  pages  that  included  the  Fissurella  descriptions  are 
correctly  dated  1831.  Although  some  of  Lesson’s  types  have 
been  recognized  at  the  Paris  Museum,  P.  Bouchet  reports 
(personal  communication)  that  he  has  been  unable  to  locate 
any  of  the  Fissurella  types. 

By  1831,  six  of  the  currently  recognized  species  had  been 
established.  Eleven  names  had  then  been  proposed,  but  very 
few  of  these  taxa  had  been  illustrated. 

In  1835,  no  less  than  13  names  were  introduced  by  G.B. 
Sowerby,  based  on  specimens  received  from  the  British  col- 
lector Hugh  Cuming,  who  had  lived  in  Valparaiso,  Chile, 
from  1 8 1 9 to  1831.  Brief  descriptions  were  given  in  the  Pro- 
ceedings of  the  Zoological  Society  of  London  for  1 834.  Dating 
for  these  species  has  frequently  been  cited  as  1834,  but  the 
publication  date  for  the  pages  involved  is  1835.  Illustrations 
were  published  simultaneously  in  the  “Conchological  Illus- 
trations” (Sowerby,  1 835b).  Sowerby  recognized  some  of  the 
species  described  earlier  by  French  authors;  seven  of  his  names 
remain  useful:  F.  maxima,  F.  latimarginata,  F.  limbata,  F. 
oriens,  F.  pulchra,  F.  lata  (here  F.  picta  lata),  and  F.  fulves- 
cens  (here  F.  oriens  fulvescens).  The  Sowerby  types  are  pre- 
served at  the  British  Museum  (Natural  History). 

By  1835,  1 1 of  the  13  species  I recognize  from  Chile  had 
been  named,  and  the  total  number  of  described  taxa  had 
reached  25. 

R.A.  Philippi  (1845,  1845-46)  proposed  five  names,  but 


none  of  these  has  any  current  utility.  One  of  these,  F.  alba, 
has  been  used  by  some  authors,  but  is  here  regarded  as  a 
synonym  of  F.  oriens.  Unfortunately,  the  present  where- 
abouts of  Philippi’s  Fissurella  types  is  unknown. 

A. A.  Gould  (1846)  introduced  one  name,  here  regarded  as 
a synonym  for  F.  peruviana.  The  holotype  is  in  the  United 
States  National  Museum. 

Lovell  Reeve  (1849-50),  in  his  monograph  of  Fissurella 
in  the  Conchologica  Iconica,  added  two  more  of  the  species 
recognized  here,  F.  cumingi  and  F.  bridgesii,  both  of  which 
have  been  enigmatic  until  now.  However,  he  also  introduced 
nine  superfluous  names,  based  on  further  splitting  of  Cum- 
ing’s material.  Some  were  described  without  locality.  Reeve 
gave  colored  illustrations  for  all  the  previously  recognized 
species  but  did  not  give  any  interior  views  of  the  shells, 
thereby  not  treating  the  broad  margin,  one  of  the  most  useful 
characters.  Also,  he  did  not  always  figure  the  same  specimen 
illustrated  by  Sowerby,  a factor  contributing  to  confusion  in 
some  cases.  The  Reeve  types  are  housed  in  the  British  Mu- 
seum (Natural  History). 

An  attempt  at  summarizing  the  recognizable  species  in 
Chile  was  made  by  L.H.  Hupe,  1854,  who  added  Spanish 
translations  of  original  descriptions  of  earlier  species,  and 
included  some  of  his  own  commentary,  but  gave  no  illustra- 
tions. Twenty-one  species  were  recognized.  One  new  taxon 
was  introduced,  the  renaming  of  a preoccuppied  name  of 
Philippi. 

Philippi  (1857)  proposed  another  name  now  having  no 
value.  In  1860  Philippi  briefly  treated  eight  species  of  Fis- 
surella from  Paposa  (near  Antofagasta)  in  his  “Reise  durch 
die  Wueste  Atacama  . . . .” 

G. B.  Sowerby  II’s  treatment  of  Fissurella  in  the  “Thesau- 
rus Conchyliorum”  ( 1 862)  was  scarcely  an  advance  over  that 
of  Reeve.  Specimens  illustrated  were  not  always  those  of 
Sowerby  or  Reeve.  One  additional  synonym  was  named. 
Little  new  information  was  given,  nor  was  opportunity  taken 
to  reduce  the  number  of  names.  The  figures  were  smaller 
than  those  given  by  Reeve,  and  there  were  no  interior  views. 

A.T.  de  Rochebrune  and  J.  Mabille  ( 1 885)  proposed  three 
taxa  from  the  southernmost  region,  none  of  which  were  com- 
pared to  established  species;  the  names  are  now  regarded  as 
junior  synonyms.  Two  of  the  three  type  specimens  are  housed 
in  the  Paris  Museum;  the  whereabouts  of  the  other  is  un- 
known. 

H. A.  Pilsbry’s  (1890)  treatment  of  Fissurella  in  the  “Man- 
ual of  Conchology”  was  an  admirable  effort  at  summary  and 
review.  For  most  taxa  he  provided  English  translations  of 
text  by  German  and  French  authors  and  copied  original  il- 
lustrations for  all  taxa,  whether  recognized  as  valid  or  placed 
in  synonymy.  Synonyms  were  allocated  as  far  as  possible. 
Approximately  20  were  treated  as  good  species.  However, 
the  specimens  available  to  Pilsbry  were  limited,  and  many 
questions  remained  unanswered.  Only  one  unnecessary  new 
species  was  introduced,  the  holotype  of  which  is  preserved 
at  the  Academy  of  Natural  Sciences,  Philadelphia. 

J.C.  Melvill  and  R.  Standen  (1898,  1907,  1914)  listed  and 
gave  notes  on  mollusks  from  the  Falkland  Islands.  The  Fis- 
surella species  were  briefly  treated. 


4 Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


H.  Strebel  (1907)  treated  the  Magellanic  and  Patagonian 
Fissurel/a.  For  three  of  the  species  that  I consider  to  inhabit 
this  southern  area,  he  recognized  six,  but  introduced  no  syn- 
onyms. A year  later,  Strebel  ( 1 908)  listed  F.  exquisita  from 
Paulet  Island,  Antarctic  Peninsula,  a record  that  needs  further 
verification.  Unfortunately,  the  Strebel  Collection  was  de- 
stroyed in  World  War  II  (Dance,  1966:302). 

W.H.  Dali  (1909)  listed  18  species  of  Fissurella  in  his 
checklist  of  mollusks  from  the  Peruvian  faunal  province. 
Those  pertaining  to  the  Magellanic  area  were  omitted.  Some 
of  Dali’s  records  are  now  clearly  erroneous:  F.  crassa  at  the 
Galapagos  Islands  and  both  F.  maxima  and  F.  picta  at  Man- 
ta, Ecuador.  The  records  from  Manta  had  previously  been 
cited  by  Steams  (1891). 

A.  Ziegenhom  and  H.  Thiem  (1925)  reported  upon  a col- 
lection made  in  Chile  by  L.H.  Plate.  From  a collection  of 
only  15  specimens,  they  discussed  and  illustrated  1 1 species. 
They  omitted  three  species  that  I recognize  and  treated  two 
others  as  “varieties,”  but  their  scheme  is  the  best  effort  avail- 
able in  the  literature.  Some  external  features  of  the  animal 
were  mentioned,  and  good  illustrations  of  the  shells  were 
given,  but  no  interior  views.  Among  the  subsequent  authors, 
only  Odhner  (1932)  and  Dell  (1971)  cited  their  work. 

I.  Perez-Farfante  (1943)  mentioned  only  F.  picta  in  her 
account  of  Atlantic  Fissurellidae.  She  made  this  species  the 
type  of  her  new  subgenus,  Balboaina,  which  I here  synony- 
mize  with  Fissurella,  sensu  stricto. 

The  “Catalogo  descriptivo  de  fisurelidos  Chilenos”  of  F. 
Riveros-Zuniga  (1951)  was  compiled  almost  entirely  from 
the  literature.  Text  from  previous  authors  was  translated  into 
Spanish.  Illustrations  were  single  exterior  views,  most  of  which 
were  copied  from  other  sources.  Twenty-six  species  and  three 
varieties  were  recognized.  Three  of  the  common  Peruvian 
Province  species  were  erroneously  cited  from  Fuerte  Bulnes, 
near  Punta  Arenas  in  the  Strait  of  Magellan. 

A. R.  Carcelles  (1950),  Carcelles  and  S.I.  Williamson  (195 1), 
and  Carcelles  (1953),  produced  a series  of  faunal  checklists 
for  the  Patagonian,  Magellanic,  and  Antarctic  regions,  in 
which  Fissurella  species  were  listed. 

Perez-Farfante  (1952)  proposed  the  subgenus  Carcellesia, 
with  the  new  type  species  F.  doellojuradoi,  which  I regard  as 
a synonym  of  F.  oriens.  The  subgeneric  name  is  here  regarded 
as  a synonym  of  Fissurella,  sensu  stricto. 

B.  Metivier  ( 1 969)  named  F.  tixierae  from  the  Golfo  Nue- 
vo, Argentina,  a name  here  treated  as  a geographic  subspecies 
of  F.  radiosa  Lesson.  The  type  specimen  is  preserved  at  the 
Paris  Museum. 

G.M.  Pena  (1970)  included  six  species  (five  that  I recog- 
nize) in  his  list  of  the  intertidal  mollusks  of  Peru,  and  cited 
a number  of  his  own  collecting  localities  for  each. 

R.K.  Dell  (1971)  illustrated  many  Sowerby  and  Reeve  syn- 
types  from  the  Cuming  Collection  in  the  British  Museum  in 
his  report  on  mollusks  from  the  Royal  Society  Expedition  to 
southern  Chile.  His  collection,  however,  was  not  sufficiently 
complete  to  enable  a full  revision,  and  his  list  of  1 1 “possible 
species”  differs  considerably  from  that  adopted  here. 

L.  Marincovich  (1973)  figured  the  five  most  abundant 
species  from  Iquique  in  northern  Chile  but  did  not  discuss 


their  synonymy.  The  Marincovich  collection  is  preserved  at 
the  Los  Angeles  County  Museum  of  Natural  History. 

J.  Christiaens  (1973)  did  not  treat  the  Chilean  species  in 
his  review  of  the  tropical  Fissurella  species;  however,  he 
proposed  the  subgenus  Corrina  for  F.  alba  Philippi,  a species 
here  placed  in  the  synonymy  of  F.  oriens.  Corrina  is  here 
regarded  as  a synonym  of  Fissurella,  sensu  stricto. 

J.  Ramirez-Boehme  (1974)  gave  a key  that  included  30 
different  species  (plus  two  varieties)  of  Fissurella  from  Chile. 
Some  of  the  taxa  recognized  in  his  key  have  never  been 
illustrated  and  have  been  considered  indeterminate  by  other 
authors.  He  also  introduced  four  synonymous  names  in  Fis- 
surella (along  with  2 1 “new  species”  names  for  acmaeid  lim- 
pets). All  were  figured  in  watercolor.  Types  are  preserved  at 
the  Museo  Nacional  de  Historia  Natural,  Santiago. 

In  July,  1978,  I distributed  copies  of  a preliminary  draft 
of  this  manuscript  to  a number  of  Chilean  biologists  whom 
I had  met  in  1975.  That  version  differed  from  this  primarily 
in  recommending  the  replacement  of  F.  rudis  Deshayes,  1 830, 
for  F.  costata  Lesson,  1831.  However,  Deshayes’  name  is 
preoccupied,  so  the  better  known  name  of  Lesson  is  rein- 
stated here.  I am  gratified  to  see  that  my  present  classification 
scheme  has  been  adopted  by  Chilean  biologists,  and  am  grate- 
ful for  the  help  they  have  provided  in  making  this  account 
the  more  complete. 

Recent  papers  on  the  biology  of  Fissurella  species  are  those 
ofAcuna(1977),  Bretos(1978,  1979,  1980,  1982,  1983),  Jara 
and  Moreno  (1984),  Moreno  and  Jaramillo  (1983),  and  Mo- 
reno et  al.  (1984). 

STRUCTURE 
Internal  Anatomy 

Anatomy  in  the  Lissurellidae  has  been  treated  by  Boutan 
(1885),  Illingworth  (1902),  Tobler  (1902),  Ziegenhorn  and 
Thiem  (1925),  and  Odhner  (1932).  The  latter  two  accounts 
included  references  to  Chilean  species  of  Fissurella.  Lretter 
and  Graham  (1962)  gave  a number  of  useful  drawings  of 
fissurellid  anatomy.  The  reader  is  referred  to  these  works  for 
details. 

Although  some  incidences  of  hermaphroditism  are  known 
in  fissurellids  (see  Lretter  and  Graham,  1964),  to  my  knowl- 
edge, Fissurella  species  are  gonochoristic,  having  separate 
sexes.  There  are  no  apparent  external  sexually  dimorphic 
features,  although  the  testis  of  males  is  beige-colored  and  the 
ovary  of  females  is  bright  green.  The  gonads  discharge  through 
the  right  kidney,  which  therefore  has  a reproductive  as  well 
as  an  excretory  function.  The  Lissurellidae  are  unusual  among 
archaeogastropods  in  having  a highly  reduced  left  kidney, 
which  is  nearly  vestigial.  In  contrast,  the  left  kidney  is  a 
prominent  papillary  sac  in  the  archaeogastropod  families 
Pleurotomariidae,  Haliotidae,  and  Trochidae.  These  families 
also  differ  in  having  the  spiral  caecum  appendage  to  the 
stomach  and  a well-developed  hypobranchial  gland  attached 
to  the  mantle  skirt,  structures  that  are  lacking  in  the  Lissu- 
rellidae. 

Anatomy  in  the  Lissurellidae  is  so  unlike  that  of  the  Pleu- 
rotomariidae, Haliotidae,  and  Trochidae,  that  the  affinity  is 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  5 


Figure  2.  External  anatomy  of  Fissurella  picta.  Two  views  of  same 
preserved  specimen  removed  from  shell,  the  mantle  skirt  cut  above 
the  head  to  the  excurrent  siphon  and  folded  back  to  show  the  paired 
gills  in  the  mantle  cavity.  Bahia  York,  Isla  de  los  Estados,  Argentina. 
LACM  71-177,  shell  length  75.4  mm.  Left,  anterior  view,  showing 
radular  ribbon  protruding  from  mouth.  Right,  dorsal  view. 


distant  from  these  groups.  I have  argued  (McLean,  1984a) 
in  support  of  the  theory  that  the  Fissurellidae  were  derived 
from  the  extinct  Paleozoic  Bellerophontacea.  The  bilateral 
symmetry  of  the  Fissurellidae  would  therefore  not  be  sec- 
ondarily derived,  as  has  been  assumed  by  most  authors,  but 
primitive. 

External  Anatomy 

Structures  of  the  head  are  the  snout,  which  terminates  in  a 
broad  oral  disc  (Fig.  2),  and  cephalic  tentacles,  the  eyes  at 
the  bases  of  the  tentacles.  The  cephalic  tentacles  extend  for- 
ward when  the  animal  is  active. 

The  body  is  attached  to  the  shell  by  a horseshoe-shaped 
shell  muscle,  which  is  open  anteriorly,  corresponding  to  the 
mantle  cavity  above  the  head.  The  animal  may  be  detached 
from  the  shell  by  severing  the  shell  muscle.  Structures  within 
the  mantle  cavity  (Fig.  2)  may  then  be  observed  by  cutting 
the  thin  mantle  roof  tissue  above  the  head.  This  exposes  a 
pair  of  large,  bipectinate  gills,  which  fill  most  of  the  space  in 
the  mantle  cavity  on  either  side.  The  gills  are  attached  by  a 
long  ventral  (efferent)  membrane  and  a short  dorsal  (afferent) 
membrane.  The  anus  opens  close  to  the  foramen.  Water  cur- 
rents enter  above  the  head,  pass  the  gills,  and  exit  through 
the  foramen  of  the  shell,  sweeping  the  fecal  material  out  at 
the  same  time.  The  currents  are  propelled  by  bands  of  cilia 
on  the  gill  filaments.  Left  and  right  kidney  openings  are  po- 
sitioned near  the  anus. 

In  living  Fissurella , the  shell  edge  is  enveloped  by  the 
mantle  fold,  which  secretes  and  protects  the  growing  edge  of 


Figure  3.  Fissurella  oriens,  SEM  view  of  juvenile  shell  1.9  mm  in 
length,  showing  coiled  protoconch  and  early  foramen.  Islota  Nihuel, 
Chiloe  Province,  Chile.  LACM  75-42. 


the  shell  and  has  sensory  papillae  scattered  on  its  surface. 
The  mantle  fold  is  color-banded  to  match  the  rayed  pattern 
of  the  shell.  The  mantle  fold  is  capable  of  expanding  to  cover 
the  entire  shell  and  foot  sides.  The  foramen  is  also  bordered 
by  mantle  folds  that  have  papillae  and  a pigment  pattern 
similar  to  that  of  the  mantle  at  the  shell  margin.  The  diameter 
of  the  excurrent  opening  in  the  mantle  skirt  varies  under 
differing  conditions  of  exposure  to  air  or  water. 

The  pigmented  side  of  the  foot  has  a single  row  of  short, 
stubby  epipodial  tentacles,  extending  anteriorly  to  the  head. 
These  tentacles  are  poorly  developed.  The  elaborate  mantle 
lobes  probably  have  more  of  a sensory  function  than  do  the 
epipodial  tentacles.  In  other  archaeogastropod  families,  par- 
ticularly the  Haliotidae,  the  epipodium  is  well  developed, 
forming  several  rows  of  tentacles  on  a separate  fold  called 
the  epipodial  lobe. 

Radula 

The  radula  consists  of  rows  of  chitinous  teeth  on  a long  ribbon 
(shown  projecting  through  the  mouth  in  Fig.  2).  The  entire 
radular  ribbon  may  be  as  much  as  '/j  the  length  of  the  shell. 
The  teeth  rows  are  rhipidoglossate,  with  a narrow  rachidian 
(central  tooth),  four  pairs  of  lateral  teeth  shaped  like  the 
rachidian,  a pair  of  large,  four-cusped  outer  lateral  teeth,  a 
pair  of  uncusped  lateromarginal  plates,  and  a large  number 
of  marginal  teeth.  The  lateromarginal  plates  separate  the  large 
outer  laterals  from  the  “books”  of  marginal  teeth.  The  fis- 
surellid  radula  is  markedly  asymmetrical  (Hickman,  1981, 
1984),  with  teeth  on  the  left  side  of  the  ribbon  higher  or  more 
anteriorly  placed  than  those  on  the  right,  extending  forward 
of  the  rachidian,  which  itself  is  asymmetrical.  This  pro- 
nounced asymmetry  enables  the  large  outer  laterals  to  inter- 
lock like  the  teeth  of  a zipper  when  the  ribbon  is  folded  and 
retracted  at  the  close  of  the  feeding  stroke.  The  teeth  are 
folded  in  the  same  way  while  developing  in  the  radular  sac. 


6 Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


Figures  4 through  7.  Shell  structure  of  Fissurella  latimarginata.  SEM  views  of  single  fragment  from  young  specimen,  courtesy  H.A.  Low- 
enstam.  LACM  64-16,  Iquique,  Chile.  (4)  Interior  view  of  shell  fragment,  the  broad,  beveled  interior  margin  (calcitic  layer  of  exterior)  of 
shell  at  right  (length  at  margin  2.14  mm),  and  the  smooth  interior  aragonitic  layer  in  center  and  left.  Fractured  area  at  lower  left  exposes  the 
crossed  lamellar  structure  of  the  aragonitic  layer  and  the  platy  calcitic  layer  below,  x 30.  (5)  Enlargement  of  lower  left  area  of  shell  fragment, 
showing  the  smooth  interior  at  top,  the  fractured  surface  of  crossed  lamellar  aragonite  below,  and  the  transition  between  the  latter  and  the 
fractured  platy  calcitic  layer  at  the  lower  edge  of  the  frame,  x 200.  (6)  Enlargement  of  upper  left  corner  of  shell  fragment.  Vertical  lines  are 
the  lamellae  of  the  smooth  interior  aragonitic  layer.  Curved  lines  represent  the  successive  positions  of  the  expanding  muscle  attachment  area. 
xl20.  (7)  Enlargement  of  lower  right  comer  of  fragment,  showing  the  smooth  growing  edge  (calcitic  layer),  the  undulations  reflecting  the 
external  sculpture  of  radial  ribs.  The  broad,  beveled  margin  has  an  irregular  surface  (for  mantle  contact)  between  the  smooth  edge  and  the 
smooth  aragonitic  surface  at  the  left,  x 80. 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  1 


The  large  outer  laterals  are  the  functional  teeth;  the  rachidian 
and  inner  laterals  are  so  small  that  their  role  is  minimal. 

Shell  Morphology 

The  apical  perforation,  or  foramen,  is  the  most  striking  shell 
feature.  It  enlarges  as  the  shell  grows;  shell  material  is  re- 
sorbed  by  the  mantle  tissue  that  surrounds  the  foramen.  Young 
shells  have  a coiled  protoconch,  but  this  is  obliterated  by  the 
expanding  foramen  and  is  generally  present  only  in  juvenile 
specimens  of  up  to  two  mm  in  length  (Fig.  3).  Although  the 
postprotoconch  stage  of  Diodora  has  a selenizone  (slitband), 
this  is  lacking  altogether  in  developing  stages  of  Fissurella 
(McLean,  1984a). 

Prominent  features  of  the  shell  interior  are  the  horseshoe- 
shaped muscle  scar,  open  in  front,  and  the  apical  callus,  a 
broad  flat  area  surrounding  the  foramen.  The  two  round 
terminations  of  the  muscle  scar  are  connected  by  a line  that 
marks  the  anterior  attachment  zone  of  the  mantle  skirt. 

Shell  Structure 

Most  fissurellids  have  the  shell  composed  entirely  of  the 
aragonitic  form  of  calcium  carbonate  (Boggild,  1930; 
MacClintock,  1963,  1967).  Boggild  (1930)  noted  that  the 
Chilean  Fissurella  crassa  also  has  an  outer  layer  composed 
of  the  calcitic  form  of  calcium  carbonate.  Other  species  treat- 
ed here  were  not  mentioned  by  Boggild,  but  all  have  a similar 
two-layered  shell  (Figs.  4-7).  The  shell  structure  of  radial  ribs 
and  the  pigmentation  is  confined  to  the  outer  layer.  This 
layer  has  a waxen,  translucent  appearance.  A periostracum 
is  lacking. 

In  the  Peruvian  and  Magellanic  species,  it  is  the  outer 
calcitic  layer  that  comprises  the  broad,  pigmented  interior 
margin  of  the  shell.  This  layer  is  secreted  by  the  mantle  lobe 
only  at  the  growing  edge.  In  contrast,  the  opaque  white  ara- 
gonitic inner  layer  is  deposited  throughout  the  interior  and 
thickens  with  growth.  The  interior  aragonitic  layer  is  thick 
only  in  the  apical  region,  where  it  has  greatly  augmented  the 
thin  calcitic  layer  of  the  early  shell.  Away  from  the  apical 
area,  the  inner  layer  becomes  thinner,  its  depth  only  one- 
third  to  one-fifth  the  thickness  of  the  calcitic  layer;  it  is  lacking 
altogether  at  the  shell  edge  (margin). 

The  prismatic  structure  of  the  calcitic  layer  is  not  readily 
apparent  under  low  magnification;  however,  the  structure  of 
the  opaque  white  aragonitic  layer  can  be  seen  under  the  dis- 
secting microscope.  It  shows  a series  of  lines  running  parallel 
to  the  shell  margin,  a typical  feature  of  “concentric  cross- 
lamellar”  shell  structure.  The  lamellae  may  be  seen  through- 
out the  interior  of  the  shell,  including  the  muscle  scar  and 
the  apical  callus  (Fig.  6). 

Tropical  species  of  Fissurella  have  shells  composed  en- 
tirely of  aragonite  and  lack  the  distinctively  colored  inner 
shell  margin.  This  difference  between  the  tropical  species 
(Fig.  30)  and  the  cooler-water  species  is  here  treated  as  a 
subgeneric  distinction. 

In  molluscan  species  with  both  calcite  and  aragonite  de- 
posited in  separate  shell  layers,  the  ratio  of  calcite  to  aragonite 
deposition  varies  with  temperature.  A greater  percentage  of 


calcite  deposition  takes  place  at  colder  latitudes  and  season- 
ally, during  winter  months  (Lowenstam,  1954,  1964;  Ver- 
meij,  1978).  Lowenstam  (1954)  noted  a greater  percentage 
of  calcite  deposition  in  species  of  Mytilus  and  Littorina  as 
latitude  increased.  This  is  apparent  in  specimens  of  Fissurella 
picta  from  different  latitudes.  In  F.  picta  from  the  Strait  of 
Magellan  at  53°37'  S (Fig.  145)  the  aragonitic  layer  is  no- 
ticeably thinner  than  in  F.  picta  from  42°42'  S (Fig.  144). 
The  greater  calcitic  deposition  in  cold  water  helps  to  explain 
why  these  species  are  so  prolific  at  high  latitudes,  where  they 
reach  a much  larger  size  than  do  their  tropical  counterparts. 
The  calcitic  layer  of  the  South  American  species  is  evidently 
an  adaptation  to  cold  water. 

Calcite  is  more  stable  than  aragonite,  and  fossil  calcitic 
shells  are  generally  better  preserved.  This  is  evident  in  the 
specimen  of  the  Pliocene  F.  concolor  Philippi,  1887,  from 
Antofagasta  (Fig.  17).  Only  the  calcitic  outer  layer  remains; 
the  aragonitic  interior  is  completely  missing.  However,  spec- 
imens in  old  shell  piles  indicate  that  when  exposed  to  sub- 
aerial weathering,  the  calcitic  layer  fractures  and  separates, 
whereas  the  aragonitic  layer  tends  to  remain  intact. 

BIOLOGY  AND  ECOLOGY 

Habitat 

As  in  other  limpet  families  with  large  numbers  of  sympatric 
species,  each  of  the  Peruvian-Magellanic  Fissurella  species 
has  a unique  habitat  or  niche. 

All  tropical  species  of  Fissurella  are  limited  to  the  intertidal 
zone,  but  some  of  the  Peruvian-Magellanic  species  extend 
into  the  subtidal  zone.  Fissurella  peruviana,  F.  pulchra,  and 
F.  oriens  may  occur  more  abundantly  in  the  sublittoral  than 
in  the  lower  intertidal  zone.  Fissurella  maxima,  F.  cumingi, 
and  F.  latimarginata  occur  commonly  from  the  lower  inter- 
tidal zone  to  a depth  of  about  5 m.  The  intertidal  occurrence 
of  these  species  is  limited  to  areas  protected  from  strong  wave 
exposure. 

The  remaining  species  are  intertidal  and  do  not  occur  in 
the  sublittoral  zone.  The  highest  occurring  species  is  Fissu- 
rella crassa,  which  is  tightly  wedged  in  crevices  when  exposed 
at  low  tide.  Fissurella  limbata  and  F.  costata  live  exposed 
to  surf  in  the  lower  intertidal  zone,  F.  limbata  on  horizontal 
surfaces,  and  F.  costata  on  vertical  surfaces.  Fissurella  nigra 
occurs  on  the  undersides  of  large  rocks  in  protected  tide  pools 
at  mid-tidal  to  lower  intertidal  levels. 

Fissurella  bridgesii  has  a unique  habitat.  It  occurs  on  rocks 
near  sandy  areas,  unlike  the  others,  which  avoid  proximity 
to  sand. 

Fissurella  picta  has  a more  ubiquitous  occurrence.  It  is  rare 
at  its  northern  limit,  where  it  is  sympatric  with  other  species, 
but  to  the  south  of  the  southern  limit  of  most  of  the  other 
species  it  occurs  from  the  mid-tidal  to  lower  intertidal  zone 
under  various  conditions  of  exposure,  filling  niches  that  are 
occupied  by  other  species  in  the  north. 

Feeding 

Little  is  known  of  the  feeding  habits  of  Peruvian  and  Ma- 
gellanic Fissurella.  Many  genera  of  fissurellids  feed  upon 


8 Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


sponges  and  detritus  (Fretter  and  Graham,  1976),  but  Ward 
(1966a)  has  shown  that  the  tropical  species  F.  ( Cremides ) 
barbadenisis  ( Gmelin,  1791)  feeds  upon  algae.  Bretos(1978) 
indicated  that  F.  crassa  feeds  upon  such  green  algae  as  Ulva 
and  Enteromorpha. 

At  Mehuin  in  southern  Chile,  Fissurella  picta  lata  is  a 
nocturnal  herbivore,  feeding  upon  the  red  alga  Iridaea  bory- 
ana  and  the  green  alga  Ulva  rigida  (Jara  and  Moreno,  1984; 
Moreno  and  Jaramillo,  1983;  Moreno  et  al.,  1984). 

Information  on  the  diets  of  the  other  South  American 
species  is  needed. 

Reproduction  and  Growth 

Bretos  (1983)  treated  reproduction  in  F.  maxima,  finding 
that  spawning  occurred  in  late  November-December  (late 
spring)  and  again  in  July-August  (winter).  The  breeding  cycle 
of  the  tropical  F.  barbadensis  was  treated  by  Ward  (1966b). 
That  species  is  known  to  have  a pelagic  phase  of  two  to  three 
days  duration. 

Bretos  (1978,  1980)  studied  the  growth  rate  of  F.  crassa, 
finding  that  growth  is  rapid  in  early  spring  and  late  summer, 
slower  in  late  spring,  autumn,  and  winter.  Harvestable  sizes 
were  reached  in  2 to  4 years.  Two  growth  rings  were  formed 
each  year  and  were  considered  reliable  indicators  of  growth 
for  the  first  6 years,  after  which  growth  was  slowed  and  the 
rings  could  not  be  identified.  Fissurella  maxima  was  also 
determined  to  form  two  growth  rings  each  year  (Bretos,  1 982). 
Growth  in  F.  latimarginata  was  studied  by  Acuna  (1977). 

The  El  Nino  event  of  1982-1983  had  a major  effect  on 
the  Fissurella  populations  in  northern  Chile.  According  to  J. 
Tomicic  (personal  communication),  all  large  Fissurella  species 
near  shore  were  killed  during  the  austral  summer  of  1 982— 
1983.  However,  in  November,  1983,  fast  growing  juveniles 
were  reported  as  abundant. 

Epibiotic  Associations 

Fissurella  shells  offer  a surface  for  colonization  by  many 
species  of  algae  and  invertebrates.  Many  shells  are  so  en- 
crusted that  color  patterns  are  obscured  (Fig.  1).  Only  those 
species  that  occur  high  in  the  intertidal  zone  {F.  crassa ),  or 
predominantly  on  the  undersides  of  large  rocks  in  the  lower 
intertidal  (F.  nigra  and  F.  pu/chra),  have  shells  that  stay 
relatively  free  of  encrusting  organisms. 

Species  of  the  shallow  sublittoral  zone  (F.  latimarginata, 
F.  cumingi,  and  F.  maxima)  generally  have  a thick  algal  mat 
consisting  of  dense  tufts  of  finely  branched  red  algae,  which 
reaches  a height  of  5 mm  above  the  shell  (Figs.  1,  8).  This 
algal  growth  is  absent  on  the  rocky  substrate  because  grazing 
by  the  black  urchin  Tetrapygus  niger,  and  presumably  by  the 
Fissurella  species,  leaves  the  rock  barren  except  for  encrust- 
ing coralline  algae. 

Although  the  algal  mat  on  the  shells  of  a living  Fissurella 
would  be  a source  of  food  for  other  individuals  of  Fissurella 
as  well  as  the  urchins,  they  evidently  do  not  tolerate  grazing 
by  their  own  kind  or  by  the  urchins,  because  the  algal  mat 
is  usually  intact.  1 observed  thick  algal  mats  on  most  shells 
seen  in  the  course  of  diving,  those  in  shell  piles  on  the  shore. 


and  those  sold  in  markets.  The  chiton  Chaetopleura  peru- 
viana is  able  to  graze  successfully  on  the  algal  mat  of  shells 
(Fig.  8). 

Balanus psittacus  is  frequently  found  on  subtidal  Fissurella 
shells,  and  B.  flosculus  occurs  on  shells  in  exposed  intertidal 
habitats,  particularly  on  F.  costata  and  F.  limbata. 

The  mussel  Semimytilus  algosus  may  form  aggregations 
on  specimens  of  F.  latimarginata  (Fig.  74). 

Epibiotic  growths  on  Fissurella  shells  provide  a protective 
advantage,  making  it  more  difficult  for  boring  organisms  to 
penetrate  the  shell.  Those  specimens  of  F.  latimarginata  that 
have  lost  the  algal  mat  are  usually  deeply  eroded.  The  ad- 
vantage of  epibionts  to  chamid  bivalves  was  discussed  by 
Vance  (1978). 

Epibiotic  Scurria  parasitica 

Most  individuals  of  F.  crassa  and  F.  limbata,  two  species 
that  only  occur  in  the  intertidal  zone,  have  a single  (or  rarely 
two)  Scurria  parasitica,  an  acmaeid  limpet,  attached  to  “home 
scars”  on  the  shell  (Figs.  9,  10).  This  limpet  occurs  also  on 
shells  of  Scurria  viridula  and  the  chitons  Enoplochiton  niger 
and  Acanthopleura  echinata  (see  Marincovich,  1973)  and 
rarely  on  other  species  of  intertidal  Fissurella.  I am  aware  of 
no  studies  on  its  biology. 

Lindberg  (1976),  Dwyer  and  Lindberg  (1981),  and  Lind- 
berg  and  Dwyer  (1983)  described  the  home-scar  depression 
of  the  Californian  acmaeid  Collisella  scabra  on  the  shells  of 
the  mussel  Mytilus  californianus  and  the  acmaeid  limpet 
Lottia  gigantea,  noting  the  similarity  of  the  scar  to  that  formed 
by  Scurria  parasitica  on  chitons.  These  epibiotic  limpets  pro- 
duce deep  scars  on  their  host  shells  (Fig.  10),  in  which  there 
is  an  outer  depression  corresponding  to  the  shell  margin,  and 
an  inner  depression  to  correspond  to  the  breadth  of  the  foot. 
The  outer  depression  provides  a seal  that  helps  to  prevent 
desiccation  and  dislodgment.  Microscopic  examination  of 
the  home  scars  shows  the  presence  of  radular  scraping  marks, 
which  indicates  that  enlargement  of  the  scar  is  an  activity 
unassociated  with  feeding.  Lindberg  and  Dwyer  (1983)  also 
found  evidence  of  shell  dissolution  by  acidic  mucopolysac- 
charides secreted  by  the  foot  and  carbonic  anhydrase  by  the 
mantle  edge. 

The  feeding  range  of  the  limpets  studied  by  Lindberg  and 
Dwyer  (1983)  was  restricted  to  the  shells  of  the  host  mollusks. 
This  is  probably  true  for  S.  parasitica  on  shells  of  F.  limbata 
and  F.  crassa.  Scar-bearing  Fissurella  shells  do  not  have  other 
encrusting  organisms  and  are  always  deeply  eroded. 

Shell  Borers 

Cirratulid  polychaetes  of  the  genus  Dodecaceria  (identified 
both  by  K.  Fauchald  and  J.C.  Castilla)  commonly  burrow 
into  Fissurella  shells,  riddling  and  weakening  them  (Fig.  1 1 ). 
The  polychaete  tubes  open  at  the  exterior  surface,  where  they 
are  ordinarily  concealed  by  the  algal  mat.  The  inside  diameter 
of  the  tube  reaches  1.3  mm.  The  burrows  are  visible  on  the 
shell  interior,  but  do  not  break  the  surface  except  in  gerontic 
shells.  Gibson  (1978)  briefly  discussed  Chilean  species  of 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  9 


Figures  8 through  15.  Shell  epiphytes,  shell  grazers,  shell  borers.  (8)  Dense  mat  of  red  algae  on  exterior  of  F.  latimarginata,  with  chiton 
Chaetopleura  peruviana  and  grazing  trails  made  by  the  chiton.  LACM  75-29,  Los  Molles,  Aconcagua  Province,  Chile,  shell  length  106  mm. 
(9)  Epizoic  limpet  Scurria  parasitica  on  F.  crassa.  LACM  90796,  Iquique,  Chile,  shell  length  57.5  mm.  (10)  Scar  of  .S’,  parasitica,  showing 
inner  and  outer  depression  of  scar.  Same  specimen,  length  of  scar  13. 1 mm.  (11)  Shell  of  F.  latimarginata  infested  with  borings  by  cirratulid 
polychaete  Dodecaceria  sp.,  exterior  view  of  cleaned  shell  showing  openings  to  burrows;  interior  view  showing  trace  of  burrows.  LACM  75- 
29,  Los  Molles,  Aconcagua  Province,  Chile,  shell  length  115  mm.  (12)  Shell  of  F.  cumingi  removed  from  stomach  of  clingfish  Sicyases 
sanguineus,  showing  breakage  pattern  caused  by  this  predator.  LACM  75-31,  Islota  Concon,  Valparaiso  Province,  Chile,  shell  length  50.8 
mm.  (13)  Shell  of  F.  costata  with  excavated  depressions  made  by  vermetid  gastropod  Dendropoma  sp.  LACM  75-27,  beach-worn  specimen, 
Bahia  El  Teniente,  Coquimbo  Province,  Chile,  length  50  mm.  (14)  Shell  of  F.  cumingi  with  burrows  of  barnacle  Cryptophialus  minutus. 
MACN  9027-1 1,  specimen  studied  by  Tomlinson  (1969),  locality  unknown,  shell  length  83.7  mm.  (15)  Exterior  surface  of  F.  limbata  showing 
burrows  of  Cryptophialus.  LACM  75-19,  Los  Colorados,  Antofagasta  Province,  Chile,  length  of  field  21  mm. 


10  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


Dodecacaeria.  However,  the  systematics  of  the  group  re- 
mains poorly  known. 

K.  Fauchald  (personal  communication)  has  also  identified 
the  spionid  polychaete  Polydora  sp.  as  a borer  in  F.  lati- 
marginata. 

Polychaete  burrows  in  Fissure/la  shells  provide  shelter  for 
young  specimens  of  the  bivalve  Hiatella  solida,  which  may 
then  bore  further  into  the  shell,  as  has  been  reported  in  shells 
of  F.  nigra  by  Gallardo  and  Osorio  (1978). 

Acrothoracican  barnacles  (burrowing  barnacles)  penetrate 
the  shells  of  Chilean  Fissurella  species  (Figs.  14,  15).  Tom- 
linson (1969:88)  reported  the  species  Cryptophialus  minutus 
Darwin,  1854,  in  a shell  of  Fissurella  maxima.  The  specimen 
examined  by  Tomlinson  (actually  F.  cumingi)  is  illustrated 
here  (Fig.  14).  I have  found  this  pattern  of  burrows  in  a 
specimen  of  F.  limbata  from  Antofagasta  (Fig.  15)  and  in  F. 
latimarginata  from  a number  of  localities. 

Burrows  of  a vermetid  gastropod,  Dendropoma  sp.,  have 
been  noted  on  specimens  of  F.  costata  (Fig.  1 3),  forming  one- 
whorled  depressions  nearly  flush  with  the  surface  of  the  host 
shell,  but  not  penetrating  to  the  interior. 

Parasites 

Bretos  and  Jiron  (1980)  reported  that  digenetic  trematodes 
of  the  genus  Proctoeces  Odhner,  1911,  family  Fellodisto- 
midae,  were  present  in  the  gonads  of  eight  species  of  Fis- 
surella in  northern  Chile.  Percentages  of  infected  individuals 
in  each  species  ranged  from  14%  to  97%.  The  effect  of  this 
parasitism  on  reproduction  in  the  host  species  is  unknown. 


Seastar  Predators 

The  seastar  He/iaster  helianthus  is  a voracious  predator  upon 
many  species  of  mollusks  in  the  lower  intertidal  zone  (Paine 
and  Palmer,  1978;  Castilla,  1981).  However,  Fissurella  has 
a highly  effective  escape  response.  Fissurella  respond  to  ini- 
tial contact  with  Heliaster  by  first  raising  the  mantle  fold 
above  the  edge  of  the  shell,  preventing  the  seastar’s  tube  feet 
from  making  contact  with  the  shell;  thereupon  they  move 
rapidly  out  of  reach.  A collector  with  a seastar  in  hand  may 
dislodge  tightly  wedged  specimens  without  using  a tool.  The 
raising  of  the  mantle  to  cover  most  of  the  shell  surface  is 
similar  to  the  response  of  the  north  Pacific  fissurellid  Diodora 
aspera  to  various  seastar  predators  (Margolin,  1964). 

Fissurella  costata  is  the  only  species  that  fails  to  show  an 
escape  response  to  Heliaster.  It  remains  tightly  appressed. 
Its  foramen,  the  smallest  among  the  larger  species,  is  evi- 
dently too  small  for  penetration  by  Heliaster. 

The  overall  effect  of  Heliaster  on  populations  of  various 
species  of  Fissurella  probably  is  not  significant.  Large  indi- 
viduals can  move  fast  enough  to  escape  and  the  small,  less 
motile  ones  stay  out  of  reach  in  crevices,  or  on  the  undersides 
of  rocks. 

In  southern  Chile,  the  asteroid  Meyenaster  gelatinosus  is 
a major  predator  on  many  mollusks  (Dayton  et  al.,  1977). 
Fissurella  and  other  gastropods  escape  predation  from  this 


seastar  in  surging  water  by  allowing  the  water  motion  to  move 
them  away. 

Vertebrate  Predators 

Non-human  vertebrate  predators  that  include  Fissurella 
species  in  their  diets  in  central  Chile  are  the  Chilean  sea  otter 
Lutra  felina,  the  seagull  Larus  dominicanus,  the  oyster  catch- 
er Haematopus  ater,  and  the  clingfish  Sicyases  sanguineus 
(see  Castilla,  1981). 

The  Chilean  sea  otter,  the  “nutria  de  mar,”  has  a restricted 
and  localized  distribution,  but  where  it  occurs,  the  effect  of 
this  carnivore  is  significant.  It  feeds  upon  Sicyases,  Concho- 
lepas,  Fissurella  species,  the  acmaeid  limpets  Scurria  species 
and  at  least  three  species  of  crabs.  Castilla  and  Bahamonde 
(1979)  gave  a more  complete  account  of  the  ecology  of  Lutra 
felina. 

According  to  Castilla  (1981),  Haematopus  ater  feeds  upon 
Concholepas,  Scurria  species  and  Fissurella  species;  Larus 
dominicanus  feeds  upon  Concholepas,  crabs,  herbivorous 
snails,  Fissurella  species,  chitons,  Scurria  species,  and  mus- 
sels. 

Sicyases  feeds  on  a wide  variety  of  invertebrates  and  algae 
on  vertical  walls  in  the  surf-exposed  intertidal  zone  (Paine 
and  Palmer,  1978).  Those  authors  reported  small  specimens 
of  several  species  of  Fissurella,  and  even  one  relatively  large 
specimen  of  F.  cumingi  (Fig.  12),  in  clingfish  stomachs.  Most 
of  the  Fissurella  shells  were  broken  at  one  end,  presumably 
by  the  strong  teeth  of  this  predator.  Many  shells  cast  up  on 
beaches  are  broken  in  a similar  way,  suggesting  that  Sicyases 
is  a major  predator  on  Fissurella.  A study  of  the  breakage 
pattern  in  beach-worn  shells  would  be  useful  to  further  doc- 
ument the  feeding  of  Sicyases. 

Fissurella  costata  is  well  adapted  to  habitats  where  Si- 
cyases occurs.  It  attaches  tightly,  making  it  difficult  for  the 
clingfish  to  get  hold  of  the  shell.  Other  species  of  Fissurella 
have  poor  defense  against  Sicyases  because  the  shell  edge 
normally  is  raised  and  the  mantle  and  foot  exposed.  Sicyases 
may  be  such  an  effective  predator  that  it  completely  removes 
other  species  that  stray  into  its  habitat. 

Human  Predation  and  Economic  Importance 

Man  is  the  chief  predator  upon  Fissurella.  Large  individuals 
of  all  species  are  used  for  food  throughout  Chile  and  Peru.  I 
found  six  species  for  sale  in  the  municipal  market  at  Iquique: 
F.  crassa,  F.  maxima,  F.  latimarginata,  F.  cumingi,  F.  lim- 
bata, and  F.  bridgesii.  They  are  collectively  know  as  “lapas” 
and  are  not  sorted  by  species  when  sold.  Although  the  fishery 
for  the  lapa  is  on  a small  scale  compared  to  that  of  the  “loco,” 
Concholepas,  it  amounts  to  a significant  predation  pressure 
on  the  larger-shelled  species.  Those  sold  in  the  market  are 
kept  intact  in  the  shell.  However,  the  shorelines  in  Chile  have 
numerous  piles  of  discarded  shells  (Fig.  1).  According  to 
figures  from  the  Chilean  Servicio  Nacional  de  Pesca  (SER- 
NAP),  451,000  tons  of  Fissurella  species  were  harvested  in 
1982  (C.A.  Moreno,  personal  communication). 

In  populated  areas,  human  predation  on  Fissurella  is  sig- 
nificant. Moreno  et  al.  (1984)  found  few  specimens  of  F. 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  1 1 


picta  lata  over  4 cm  in  length  in  the  vicinity  of  Valdivia. 
Much  larger  sizes  occurred  in  areas  where  human  access  was 
restricted.  Where  the  Fissurella  were  experimentally  re- 
moved from  the  habitat,  there  were  dramatic  increases  in  the 
algal  cover  of  Iridaea  boryana,  its  chief  food  source.  This 
alga  is  also  harvested  in  Chile.  Human  predation  on  Fissu- 
rella therefore  helps  to  ensure  a good  harvest  of  the  alga. 

Fishermen  in  Chile  know  each  species  by  a common,  de- 
scriptive name  (Bretos,  personal  communication).  The  names 
in  use  in  northern  Chile  are  mentioned  in  the  species  ac- 
counts. 

SYSTEMATIC  CHARACTERS 

Useful  shell  characters  include:  size;  outline  in  dorsal  view; 
sculpture— the  strength  and  spacing  of  the  radial  ribs;  color 
pattern;  interior  shell  margin  — the  thickness  and  pigmenta- 
tion pattern;  and  foramen— the  size,  shape,  and  placement. 
Other  shell  characters  such  as  the  muscle  scar  and  the  internal 
callus  show  few  significant  differences  and  are  therefore  not 
treated  unless  they  have  unusual  features.  The  organisms  that 
encrust  the  surface  of  shells  can  provide  important  ecological 
information,  but  shells  must  be  cleaned  in  order  to  see  the 
sculpture  and  color  pattern. 

Juvenile  shells  are  commonly  very  different  from  mature 
shells  and  are  therefore  separately  described  in  this  account. 

Features  of  external  anatomy  such  as  color  of  the  cephalic 
tentacles  and  development  of  mantle  lobe  papillae  and  tu- 
bercles on  the  foot  are  specific  characters  that  may  enable 
identification  of  species. 

The  morphology  of  the  large  outer  lateral  tooth  of  the 
radula  provides  a specific  character. 

The  most  useful  specific  characters  are  discussed  in  greater 
detail  as  follows. 

Size  and  Shape 

The  anterior  end  of  the  shell  is  narrower  than  the  posterior. 
The  muscle  scar  opens  anteriorly.  All  shells  are  illustrated 
here  with  the  anterior  at  the  top;  the  lateral  view  shows  the 
left  side. 

The  outline  in  most  species  is  elongate-oval.  Fissurella 
costata,  F.  picta  lata,  and  F.  peruviana  are  generally  rounder 
than  other  species.  Fissurella  latimarginata,  F.  cumingi,  and 
F.  pulchra  are  wedge-shaped,  having  a relatively  narrow  front 
end  and  tapered  sides. 

Shells  seldom  lie  flat  in  one  plane.  In  most  species,  the 
sides  are  slightly  elevated  relative  to  the  ends.  This  is  par- 
ticularly true  of  such  highly  motile  species  as  F.  maxima, 
and  enables  a better  fit  on  rounded  rock  surfaces.  Species 
that  commonly  nestle  in  crevices  or  have  a habitual  site  of 
home  attachment  may  instead  have  elevated  ends  for  a better 
fit.  Both  extremes  are  possible  in  F.  nigra  and  F.  oriens.  Some 
specimens  of  these  species  have  both  elevated  sides  and  el- 
evated ends,  so  that  the  shell  rests  on  four  comers. 

Shell  height  is  fairly  constant  in  some  species  and  variable 
in  others.  Species  with  relatively  low  shells  (length  3.2  to  5.9 
times  height)  include  F.  crassa,  F.  bridgesii,  and  F.  pulchra. 
Those  that  vary  from  low  to  medium  in  height  (length  2.8 

12  Contributions  in  Science,  Number  354 


to  4.5  times  height)  include  F.  maxima,  F.  latimarginata,  F. 
costata,  F.  radiosa,  and  F.  oriens.  Fissurella  peruviana  varies 
from  extremely  low  to  high  (length  1.5  to  4.8  times  height). 

In  some  species,  the  shell  may  be  steeply  conical  in  young 
stages  and  abruptly  become  more  flattened  at  later  stages. 
This  commonly  happens  in  F.  oriens,  F.  costata,  and  F. 
limbata. 

Fissurella  nigra  may  grow  by  increasing  the  shell  height 
while  contracting  the  length  and  width,  especially  in  gerontic 
specimens.  This  makes  the  slopes  convex  and  the  shell  mar- 
gin very  thick.  This  growth  form  has  not  been  observed  in 
other  species. 

Sculpture 

Shell  sculpture  is  relatively  consistent  within  most  species. 
The  radial  ribs  produced  in  the  earliest  growth  stages  are 
called  the  primary  ribs  and  those  arising  between  the  primary 
ribs  at  later  growth  stages  are  called  the  secondary  ribs.  Sec- 
ondary ribs  attain  the  size  and  prominence  of  the  primary 
ribs  in  F.  latimarginata,  F.  cumingi,  and  F.  oriens.  Primary 
ribs  are  stronger  than  the  secondary  ribs  at  all  growth  stages 
in  F.  picta,  F.  radiosa,  F.  costata,  F.  maxima,  and  F.  peru- 
viana. Fissurella  crassa,  F.  pulchra,  and  F.  bridgesii  generally 
have  smooth  shells  in  mature  stages,  although  their  young 
stages  have  rounded  primary  ribs,  but  no  secondary  ribs. 

Color  Pattern 

Color  patterns  in  all  species  have  definite  limits  of  variation, 
extensive  in  some,  limited  in  others.  Most  species  have  a 
pattern  of  dark-colored  rays  on  a lighter  ground  color.  Least 
variable  in  color  pattern  are  F.  maxima,  F.  limbata,  and  F. 
pulchra.  Such  strongly  rayed  shells  as  those  of  F.  picta,  F. 
radiosa,  F.  cumingi,  and  F.  oriens  have  ground  colors  ranging 
from  light  to  dark  gray  or  tan  and  correspondingly  darker 
rays.  White  shells  occur  only  in  F.  oriens.  The  most  variable 
species,  having  both  rayed  forms  and  uniformly  colored  forms, 
are  F.  peruviana,  F.  latimarginata,  F.  radiosa,  and  F.  oriens. 

In  addition  to  the  radial  rays,  all  species  may  have  con- 
centric growth  bands  of  varying  color  intensity.  Changes  in 
the  coloration  of  growth  bands  have  been  correlated  with 
changes  in  diet  in  such  herbivorous  archaeogastropods  as 
Haliotis  (Olsen,  1968a,  1968b).  Changes  in  supply  of  food 
or  a shift  in  the  algal  composition  of  the  diet  can  probably 
be  correlated  with  changes  in  the  banding  of  Fissurella  shells. 
Concentric  color  changes  are  most  pronounced  in  F.  picta, 
F.  radiosa,  and  F.  oriens,  the  three  species  that  range  to  the 
high  southern  latitudes  where  ecological  conditions  are  most 
extreme.  Bretos  (1978,  1980)  has  shown  that  there  are  sea- 
sonal growth  rings  in  F.  crassa. 

Shells  exposed  to  weathering  fade.  The  dark  purple  or  gray 
rays  change  to  red,  particularly  in  F.  maxima  and  F.  picta 
lata. 

Interior  Margin 

The  interior  margin  or  border,  composed  of  the  calcitic  layer 
of  the  shell,  generally  has  several  bands  or  zones,  visible  also 

McLean:  Peruvian  and  Magellanic  Fissurella 


in  cut  or  broken  pieces  of  the  shell.  Color  differences  in  the 
margin  are  useful  specific  characters.  The  width  of  the  margin 
changes  with  growth.  In  young,  rapidly  growing  shells,  it  is 
relatively  broad;  in  mature  shells  it  is  proportionately  nar- 
rower, and  in  old  shells  it  may  be  nearly  obliterated  by  the 
encroachment  and  thickening  of  the  inner  aragonitic  layer. 
It  is  consistently  narrow  in  all  growth  stages  of  F.  radiosa 
and  F.  peruviana.  In  most  species  the  margin  or  growing  edge 
is  flat,  but  in  F.  maxima  it  is  convex,  and  in  F.  crassa  the 
entire  edge  is  rounded. 

In  some  species  the  margin  of  the  shell  is  uniformly  pig- 
mented across  its  full  width,  in  others  the  pigment  is  con- 
centrated near  the  surface  or  deeper  within  the  layer.  Only 
in  F.  nigra  is  the  outermost  zone  darker  than  the  inner  zone. 
In  F.  latimarginata  and  F.  pulchra  the  outer  edge  is  lighter 
and  in  F.  limbata  the  outer  edge  is  much  lighter  and  contrasts 
sharply  with  the  inner  zone.  Color  rays  are  confined  to  the 
outermost  layers  of  the  margin  in  F.  limbata  and  F.  maxima, 
but  extend  the  full  width  of  the  margin  in  F.  cumingi  and  F. 
oriens. 

Foramen 

The  relative  size  and  the  configuration  of  the  foramen  changes 
with  growth.  In  young  shells  it  is  elongate,  broad  in  the  mid- 
dle and  constricted  in  two  places  on  the  sides.  The  foramen 
can  be  described  as  tripartite,  and  the  side  walls  as  bidentate 
if  the  three-lobed  aspect  is  especially  conspicuous.  In  most 
species  the  foramen  changes  from  tripartite  in  young  stages 
to  oval  in  mature  stages.  In  some  species  its  size  in  mature 
specimens  varies  greatly;  it  may  become  very  large  in  some 
old  shells  of  F.  oriens.  In  F.  peruviana,  the  tripartite  aspect 
of  the  foramen  is  lost  at  a very  early  stage,  and  the  foramen 
becomes  oval.  Fissurella  costata  has  a particularly  small  fo- 
ramen at  all  growth  stages.  Fissurella  limbata  is  unusual  in 
retaining  an  elongate  foramen  in  mature  sizes.  Fissurella 
crassa  also  retains  an  elongate  foramen  that  is  constricted  in 
the  middle,  although  the  young  shells  are  bidentate  like  those 
of  other  species. 

In  most  species  the  position  of  the  foramen  is  slightly 
posterior  to  the  midpoint  of  the  shell,  but  in  F.  nigra  and  F. 
radiosa  it  is  more  markedly  so. 

Juvenile  Shell 

The  earliest  juvenile  shells  of  all  species  are  more  elevated 
and  conical  than  later  stages.  Primary  ribs  appear  at  an  early 
stage.  In  forms  with  a rayed  pattern,  the  elevated  ribs  are 
light-colored  and  the  interspaces  are  dark-rayed.  Juveniles 
of  many  species  have  a pair  of  broad  white  rays  extending 
laterally,  more  prominently  than  the  other  light-colored  rays. 
This  pattern  is  especially  evident  in  the  juveniles  of  F.  la- 
timarginata, in  which  the  light  rays  persist  until  the  shell  is 
10  or  20  mm  in  length.  In  F.  nigra,  F.  crassa,  F.  peruviana, 
F.  cumingi , and  F.  maxima,  the  two  light  rays  are  seen  only 
in  juveniles  of  less  than  5 mm  length.  Some  have  character- 
istic early  colorations  unlike  the  adults.  Young  F.  nigra  are 
light-colored  rather  than  black;  F.  oriens,  F.  maxima,  and 
F.  peruviana  are  reddish  when  young;  F.  limbata  has  a zigzag 


pattern  of  lines;  F.  maxima  and  F.  cumingi  have  speckled 
patterns. 

External  Anatomy 

The  relative  size  of  the  animal  in  proportion  to  its  shell  is  a 
useful  comparison  for  at  least  those  species  at  either  extreme; 
the  animals  of  most  species  are  relatively  large  and  just  barely 
containable  within  the  shell.  The  extremes  are  Fissurella  crassa 
and  F.  bridgesii,  which  have  flat  shells  that  cannot  contain 
the  animal,  and  Fissurella  costata  and  F.  peruviana,  which 
have  high  conical  shells,  the  animal  easily  contained  within 
the  shell. 

In  most  species,  the  cephalic  tentacles  are  dark,  reddish 
on  the  inner  side  and  yellowish  at  the  tips.  Fissurella  nigra 
is  the  only  species  that  shows  only  shades  of  gray  and  black 
on  the  tentacles  as  well  as  on  the  mantle  and  foot. 

The  mantle  lobe  has  three  edges,  here  called  the  inner,  the 
upper,  and  the  lower.  The  inner  lobe  lacks  papillae  and  is  in 
direct  contact  with  the  growing  margin  of  the  shell.  The  upper 
lobe  extends  up  over  the  edge  of  the  shell,  and  the  lower  lobe 
extends  down.  The  edges  of  the  upper  and  lower  lobes  have 
finely  branched  papillae.  The  papillae  of  the  upper  lobe  are 
generally  more  strongly  developed  than  those  of  the  lower 
edge.  The  area  between,  which  is  greatly  expandable,  is  usu- 
ally vertically  banded  to  match  the  pattern  of  rays  on  the 
shell.  This  area  may  also  show  dark  pigment  in  concentric 
grooves.  The  edge  of  the  lower  lobe  of  F.  latimarginata  is  a 
striking  orange  color,  the  only  species  so  marked,  making  it 
readily  recognizable. 

In  all  species  the  side  of  the  foot  is  rugose  or  pustular. 
Coloration  is  mottled,  the  tips  of  the  pustules  or  tubercles 
lighter  in  color.  Overall  coloration  of  the  foot  is  brown  or 
gray  in  most  species;  however,  the  foot  of  F.  cumingi  has  a 
distinctive  strawberry-red  color  and  that  of  F.  costata  has  a 
pale  pinkish-brown  color. 

Epipodial  tentacles  extend  along  the  foot  sides.  They  are 
short  and  stubby  but  are  slightly  more  prominent  that  the 
ordinary  tubercles  on  the  foot  side.  They  are  particularly 
prominent  in  F.  oriens. 

Radula 

There  are  few  specific  differences  in  the  rachidian  and  inner 
lateral  teeth  in  Fissurella,  although  those  of  F.  pulchra  (Figs. 
266,  267)  are  somewhat  unusual  in  having  longer  overhang- 
ing cusps.  The  larger  outer  laterals,  however,  show  interspe- 
cific differences,  as  will  be  noted  in  comparing  the  illustra- 
tions for  the  radula  of  F.  nimbosa  (Figs.  21,  25),  F.  picta 
(Figs.  138,  142,  143),  F.  peruviana  (Figs.  43,  44),  F.  oriens 
(Figs.  193,  194),  and  F.  pulchra  (Figs.  266,  267). 

NAMES,  ALLOCATIONS,  AND 
KEY  CHARACTERS 

Names  and  Allocations 

Fifty-eight  names  have  been  proposed  for  Recent  species  of 
Fissurella  from  the  Peruvian  Faunal  Province,  which  encom- 
passes central  Peru  to  central  Chile,  and  the  Magellanic  Fau- 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  13 


Table  1.  Names  proposed  for  Recent  species  of  Fissurella  from  the 
Peruvian  and  Magellanic  faunal  provinces,  with  allocations  as  dis- 
cussed in  this  paper. 

affinis  Sowerby,  1835  = F.  peruviana 

alba  Philippi,  1845  = F.  oriens  oriens 

arenicola  Rochebrune  and  Mabille,  1885  = F.  oriens  oriens 

atrata  Reeve,  1850  = F.  picta  picta 

australis  Philippi,  1845  = F.  oriens  oriens 

bella  Reeve,  1 849  = F.  latimarginata 

biradiata  Sowerby,  1835  = F.  latimarginata 

bridgesii  Reeve,  1 849 

clypeiformis  Sowerby,  1825  = F.  crassa 

clypeus  Sowerby,  1835  = F.  peruviana 

cheullina  Ramirez-Boehme,  1974  = F.  oriens  oriens 

chilensis  Sowerby,  1835  = F.  costata 

concinna  Philippi,  1845  = F.  maxima 

costata  Lesson,  1831 

crassa  Lamarck,  1822 

cumingi  Reeve,  1849 

darwinii  Reeve,  1 849  = F.  radiosa  radiosa 

depressa  Lamarck,  1822  = F.  crassa 

doellojuradoi  Perez-Farfante,  1952  = F.  oriens  oriens 

dozei  Rochebrune  and  Mabille,  1885  = F.  radiosa  radiosa 

exquisita  Reeve,  1850  = F.  radiosa  radiosa 

Jlavida  Philippi,  1857  = F.  oriens  oriens 

fulvescens  Sowerby,  1835  = F.  oriens  fulvescens 

galericulum  Reeve,  1 850  = F.  latimarginata 

grandis  Sowerby,  1835  = F.  nigra 

grisea  Reeve,  1 849  = F.  radiosa 

hedeia  Rochebrune  and  Mabille,  1885  = /•'.  oriens  oriens 

hondurasensis  Reeve,  1 849  = F.  maxima 

lata  Sowerby,  1835  = F.  picta  lata 

latimarginata  Sowerby,  1835 

limbata  Sowerby,  1835 

maxima  Sowerby,  1835 

mexicana  Sowerby,  1835  = F.  oriens  oriens 

multilineata,  limbata  var.,  Ziegenhom  and  Thiem,  1925  = F.  lim- 
bata 

muricata  Reeve,  1850  = F.  picta  picta 
navidensis  Ramirez-Boehme,  1974  = F.  picta  lata 
nigra  Lesson,  1831 

nigra  Philippi,  1845,  not  Lesson,  1831  = F.  radiosa  radiosa 

oblonga  Ramirez-Boehme,  1974  = F.  oriens  oriens 

obovalis  Lesson,  1831  = ? 

occidens  Gould,  1846  = F.  peruviana 

oriens  Sowerby,  1835 

papudana  Ramirez-Boehme,  1974  = F.  peruviana 
peruviana  Lamarck,  1822 
philippiana  Reeve,  1850  = F.  radiosa  radiosa 
philippii  Hupe,  1854  = F.  radiosa  radiosa 
picta  Gmelin,  1791 

polygona  Sowerby  II,  1 862  = F radiosa  radiosa 
pulchra  Sowerby,  1835 

punctatissima  Pilsbry,  1890  = F.  latimarginata 
radiosa  Lesson,  1831 

rubra,  costata  var.,  Ziegenhom  and  Thiem,  1925  = F.  costata 

rudis  Deshayes,  1830,  not  Roeding,  1798  = F.  costata 

solida  Philippi,  1845  = F.  maxima 

stellata  Reeve,  1850  = F.  cumingi 

subrotunda  Deshayes,  1830  = F.  peruviana 

tixierae  Metivier,  1969  = F.  radiosa  tixierae 

violacea  Rathke,  1833  = F.  nigra 


Figure  16.  Distribution  of  Fissurella  species  in  Peru,  Chile,  and 
southern  Argentina  by  degrees  south  latitude.  The  identified  place 
names  are  major  LACM  localities,  some  comprising  several  stations. 
The  dots  indicate  that  specimens  are  represented  in  the  LACM  col- 
lection. Lines  without  dots  indicate  distributions  taken  from  other 
sources  (see  text). 

nal  Province,  which  includes  southern  Chile  and  southern 
Argentina.  These  names  are  listed  alphabetically  in  Table  1, 
followed  by  my  allocation.  In  the  text  that  follows,  I discuss 
the  limits  of  variation  for  each  species,  but  do  not  use  variety 
or  form  names,  even  though  some  may  be  based  upon  readily 
recognizable  variants.  Geographic  subspecies  are  discernible 
in  three  species  of  the  Magellanic  Faunal  Province:  F.  picta, 
F.  radiosa,  and  F.  oriens.  Trinomial  designations  are  there- 
fore used  for  these  three  species. 

Difficulty  in  determining  the  taxa  of  such  early  authors  as 
Lesson  (1831)  and  Philippi  (1845,  1857)  has  been  due  to  a 
lack  of  illlustrations.  I have  not  succeeded  in  locating  the 
type  specimens  of  these  two  authors.  Nevertheless,  using 
evidence  from  the  original  descriptions  and  type  localities,  I 
feel  confident  of  the  allocation  of  all  names,  except  for  F. 
obovalis  Lesson,  which  is  unassigned. 

Chart  of  Key  Characters 

Attempts  to  prepare  a dichotomuous  key  have  not  been  suc- 
cessful because  of  the  extreme  variability  of  some  of  the 


14  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


species.  Instead,  the  key  characters  are  summarized  in  Ta- 
ble 2. 

DISTRIBUTION  AND  ZOOGEOGRAPHY 

Distributions  of  the  13  species  of  the  subgenus  Fissurella 
known  from  Peru,  Chile,  and  Argentina  are  shown  by  latitude 
in  Figure  16. 

Nine  species  occur  in  the  warm-temperate  Peruvian  Prov- 
ince, which  extends  from  central  Peru  to  central  Chile:  F. 
peruviana,  F.  maxima,  F.  latimarginata,  F.  cumingi,  F.  cos- 
tata,  F.  limbaia,  F.  erassa,  F.  bridgesii,  and  / '.  pulchra.  Their 
northern  and  southern  distributional  records  differ,  but  all 
nine  are  present  between  Iquique  and  Concepcion,  Chile  (20° 
S to  37°  S). 

Four  species  are  primarily  members  of  the  cold-temperate 
Magellanic  Province,  which  includes  southern  Chile  and 
southern  Argentina:  F.  picta,  F.  radiosa,  F.  oriens,  and  F. 
nigra.  Their  distributions  also  differ,  but  all  are  present  be- 
tween 43°  S and  54°  S. 

Except  for  F.  radiosa,  three  of  the  four  Magellanic  species 
extend  north  in  Chile  to  overlap  with  the  distributions  of  the 
Peruvian  species.  The  region  of  overlap  is  that  between  Val- 
paraiso and  Concepcion,  from  33°  S to  37°  S,  in  which  12  of 
the  1 3 species  occur.  At  some  point  to  the  south  of  Concep- 
cion, four  of  the  Peruvian  Province  species  (F.  maxima,  F. 
latimarginata,  F.  pulchra,  and  F.  bridgesii)  drop  out.  I did 
not  find  them  at  Mehuin,  Valdivia  Province  (39°  S).  Stuardo 
(1964)  noted  a transition  zone  between  the  two  provinces 
from  38°  S to  43°  S.  The  transition  zone  noted  here  is  there- 
fore considerably  to  the  north  of  that  reported  by  Stuardo. 

The  number  of  Fissurella  species  occurring  between  Val- 
paraiso and  Concepcion  is  significantly  greater  than  the  num- 
ber known  either  to  the  north  or  the  south.  Distributions  of 
species  in  other  families  should  be  considered  to  determine 
whether  this  transition  area  has  a higher  number  of  species 
than  either  of  the  two  provinces  treated  separately. 

The  northernmost  occurrence  of  the  Peruvian  Province 
species  of  Fissurella  is  at  Isla  Guanape,  Peru  (8°33'  S),  where 
I have  found  F.  latimarginata,  F.  limbata,  F.  bridgesii,  and 
F.  peruviana.  I found  none  in  January,  1974,  in  the  transi- 
tional region  between  the  Peruvian  Faunal  Province  and  the 
tropical  Panamic  Faunal  Province  at  the  Lobos  Afueras  Is- 
lands in  northern  Peru. 

None  of  the  species  is  known  from  the  offshore  islands  of 
central  Chile,  Isla  San  Felix  or  Islas  Juan  Fernandez,  either 
from  literature  records  or  recent  expedition  material  from 
ANTON  BRUUN  cruises.  A number  of  common  Chilean 
mollusks  are  known  from  these  islands.  The  absence  of  Fis- 
surella correlates  with  the  brief  planktonic  larval  stage  and 
resulting  poor  colonizing  potential  of  fissurellids  and  other 
archaeogastropod  larvae. 

Fissurella  picta,  F.  radiosa,  and  F.  oriens  are  abundant  at 
the  southern  limits  of  their  distributions,  where  they  un- 
doubtedly extend  to  Cape  Horn.  These  three  species  also 
occur  at  the  Falkland  Islands,  which  region  is  included  in  the 
Magellanic  Faunal  Province.  They  do  not,  however,  occur 
at  South  Georgia  or  any  of  the  subantarctic  islands  east  of 


the  Falklands  (Powell,  1951).  Strebel  (1908:79)  reported  F. 
exquisita  [here  = F.  radiosa]  at  Paulet  Island,  Antarctic  Pen- 
insula, a record  that  should  not  be  accepted  without  further 
confirmation. 

The  Magellanic  Faunal  Province  extends  north  through 
the  Patagonian  region  of  Argentina  to  the  Gulf  of  San  Matias, 
but  only  one  of  the  Magellanic  species,  F.  radiosa,  occurs  to 
the  north  of  Tierra  del  Fuego  in  Argentina.  It  has  a subspecies, 
F.  radiosa  tixierae,  in  its  northernmost  extent  in  Argentina. 
Absence  of  the  other  three  species  in  Chubut  and  Santa  Cruz 
Provinces  of  Argentina  may  be  due  to  the  vastly  different 
ecological  conditions.  In  Argentina,  the  tidal  range  is  ex- 
treme, broad  tidal  flats  are  exposed,  and  inshore  sediments 
often  consist  of  fine  beach  sand.  In  southern  Chile,  the  tidal 
range  is  less  extensive,  inshore  waters  are  deep  and  clear,  and 
beaches  are  few. 

FOSSIL  RECORD 

There  is  little  information  in  the  literature  about  the  fossil 
record  of  Fissurella  in  Peru  and  Chile.  Ihering  (1907)  par- 
ticularly noted  the  abundance  of  Fissurella  in  the  Recent  and 
the  complete  lack  of  the  genus  in  the  lower  Tertiary  of  Chile 
and  Patagonia. 

In  the  most  recent  report  on  the  mollusks  of  the  Pliocene 
and  Pleistocene  formations  of  Chile  (Herm,  1969),  none  of 
the  Fissurella  species  was  given  formal  systematic  treatment. 
Herm  listed  five  characteristic  Pleistocene  species:  F.  micro- 
trema  Sowerby,  1 833  [undoubtedly  F.  peruviana,  rather  than 
the  tropical  F.  microtrema],  F.  costata,  F.  erassa,  F.  lata  [F. 
picta  lata  here],  and  F.  concinna  [F.  maxima  here].  In  his 
list  of  Pliocene  species  he  noted  only:  “Fissurella,  div.  sp.” 

One  species  has  been  described  from  fossil  material:  F. 
concolor  Philippi,  1887,  from  Pliocene  beds  of  Mejillones, 
north  of  Antofagasta,  Chile.  I have  received  specimens  iden- 
tified as  this  species  from  E.  Martinez,  collected  from  two  of 
the  Pliocene  localities  near  Antofagasta  detailed  by  Herm: 
the  Cerro  Costino  locality  south  of  Antofagasta,  and  the  Hor- 
nito  locality  north  of  Antofagasta.  Specimens  (Fig.  1 7)  agree 
with  Philippi’s  description  in  having  three  smaller  ribs  be- 
tween each  of  the  larger  ribs.  The  overall  shape  and  profile 
is  similar  to  that  of  F.  maxima,  but  the  primary  ribs  are 
stronger  than  those  of  F.  maxima.  The  interior  aragonitic 
layer  is  missing  entirely  from  the  specimens,  in  agreement 
with  the  principle  that  calcitic  structures  are  best  preserved 
in  fossils. 

A fragmentary  specimen  of  another  species  from  the  Cerro 
Costino  locality  has  also  been  received  from  E.  Martinez. 
This  species  (Fig.  18)  has  some  affinity  to  F.  erassa.  In  the 
absence  of  additional  specimens,  I am  unable  to  further  treat 
the  fossil  record  of  the  group. 

Fissurella,  sensu  stricto,  is  one  of  the  youngest  genera  in 
the  Fissurellidae,  traced  only  to  the  Pliocene  (Herm,  1969). 
Except  for  the  Caribbean  type  species,  it  is  an  eastern  Pacific 
genus,  well  represented  in  the  Peruvian  and  Magellanic  fau- 
nal provinces,  and  with  a single  species  ranging  from  Cali- 
fornia to  Baja  California,  Mexico  (Fig.  19).  The  type  species 
may  have  become  established  in  the  Caribbean  during  the 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  15 


I’able  2.  Chart  of  key  characters. 


Size 

Height 

Mature  sculpture 

Outline  of  base 

F.  peruviana 

small 

low  to 

medium  ribs. 

oval  to 

25-40  mm 

high 

strong  primaries 

elongate 

F.  maxima 

large 

medium 

strong  ribs. 

elongate 

80-135  mm 

strong  primaries 

oval 

F.  latimarginata 

large 

low  to 

fine  ribs 

tapered 

70-1 1 5 mm 

medium 

oval 

F.  cumingi 

large 

medium 

medium  ribs 

tapered 

80-100  mm 

oval 

F.  costata 

medium 

low  to 

strong  ribs. 

oval 

50-80  mm 

medium 

strong  primaries 

F.  picta  picta 

large 

medium 

strong  ribs. 

elongate 

65-95  mm 

strong  primaries 

oval 

F.  picta  lata 

medium 

medium 

strong  ribs. 

oval 

50-80  mm 

to  high 

strong  primanes 

F.  rad.  radiosa 

small 

low  to 

medium  nbs, 

tapered 

40-55  mm 

medium 

strong  primaries 

elongate 

F.  rad.  tixierae 

small 

medium 

medium  ribs. 

tapered 

25-45  mm 

strong  primaries 

elongate 

F.  oriens  oriens 

medium 

low  to 

fine,  broad  ribs 

elongate 

40-70  mm 

medium 

oval 

F.  o.  fulvescens 

medium 

low 

fine,  broad  ribs 

elongate 

45  mm 

oval 

F.  nigra 

large 

medium 

fine,  weak  ribs 

elongate 

70-1  10  mm 

oval 

F.  limbata 

large 

medium 

undulations 

elongate 

60-90  mm 

oval 

F.  crassa 

medium 

low 

undulations 

elongate 

60-90  mm 

oval 

F.  bridgesii 

large 

low 

irregular  striae 

tapered 

65-90  mm 

oval 

F.  pulchra 

medium 

low 

undulations 

tapered 

35-75  mm 

oval 

period  in  which  the  Central  American  seaway  provided  free 
access  between  the  western  Atlantic  and  eastern  Pacific  dur- 
ing the  Miocene  and  early  Pliocene  (see  Woodring,  1965, 
1966). 

FORMAT  FOR  SPECIES  ACCOUNTS 

Description.  Shell  descriptions  treat  the  following  char- 
acters in  order:  the  size  range  (length  in  mm)  of  examples 
considered  to  be  mature,  the  relative  height,  the  outline  in 
dorsal  view,  whether  the  sides  or  ends  are  raised,  strength  of 
the  radial  ribs,  the  color  pattern,  the  shell  layers,  the  interior 
margin,  and  the  position  and  shape  of  the  foramen.  Dimen- 
sions for  shell  length,  width,  and  height  are  given  in  that 
order  in  the  captions  for  the  figured  specimens,  not  duplicated 
in  the  text.  For  specimens  with  uneven  basal  margins,  shell 
height  is  the  maximum  elevation  when  the  shell  rests  upon 
a plane  surface. 


Juvenile  Shell.  A separate  description. 

Mantle  and  Foot.  Anatomical  characters  include  the  rel- 
ative size  of  the  body  and  shell,  the  relative  prominence  of 
papillae  on  the  upper  and  lower  edges  of  the  mantle  lobe, 
coloration  of  the  cephalic  tentacles,  the  color  of  the  foot-side 
and  the  relative  prominence  of  the  foot-side  tubercles. 

Habitat.  The  intertidal  or  subtidal  occurrence,  conditions 
of  exposure,  and  the  epibiotic  associations. 

Distribution.  The  northernmost  and  southernmost  verified 
record,  the  latitude  coordinates  for  these  records,  and  the 
source  of  the  record.  The  source  is  the  museum  catalog  num- 
ber if  the  specimen  has  been  examined,  or  an  author  and 
date,  if  the  record  is  based  on  a published  account  considered 
to  be  correctly  identified.  This  is  followed  by  commentary 
about  its  possible  occurrence  beyond  the  verified  limits  and 
corrections  of  previous  records  now  considered  to  be  inac- 
curate. 


16  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


Table  2.  Continued. 


Shell  color 

Shell  margin 

Mature  foramen 

Foot  side  color 

red,  gray,  white 

narrow  showing 

oval 

gray-brown  mottled 

rayed  or  solid 

rays 

reddish  brown 

broad,  rays  on 

oval 

dark  brown  mottled 

rays  on  white 

outer  edge 

solid  purple 

broad,  solid 

elongate 

black  with  yellow 

gray 

reddish  brown 

oval 

outer  edge  of  mantle 

reddish  rays 

broad,  rays  show 

elongate 

reddish  brown 

on  dark  yellow 

on  full  width 

oval 

mottled 

gray  rays  on 

broad,  rays  show 

elongate 

pinkish  gray 

yellow 

on  full  width 

oval,  small 

mottled 

split  gray  rays 

broad,  rays  strong 

elongate 

light  brown-black 

on  light  ground 

on  outer  edge 

mottled 

split  gray  rays 

broad,  rays  strong 

elongate 

brown-black 

on  light  ground 

on  outer  edge 

mottled 

gray  or  reddish 

narrow,  showing 

elongate 

gray-brown 

rays  on  white 

rays 

mottled 

gray  or  reddish 

narrow,  showing 

elongate 

gray-brown 

rays  on  white 

rays 

mottled 

red  to  gray  rays 

medium  width. 

elongate 

pinkish  brown 

on  light  ground 

showing  rays 

oval 

mottled 

reddish  rays  on 

medium  width. 

elongate 

not  seen 

dark  yellow  gr. 

showing  rays 

oval 

solid  gray, 

broad,  gray. 

elongate  oval 

gray  mottled 

rays  faint 

dark  at  edge 

beveled 

purple  rays  on 

broad,  purple. 

elongate 

light  gray 

dark  yellow 

white  at  edge 

mottled 

solid  brown, 

brown  upturned 

very  long, 

mantle  brown  banded 

faint  rays 

constricted 

foot  gray  mottled 

gray  brown, 

broad,  solid 

elongate 

brown-black 

rays  faint 

reddish  brown 

oval 

mottled 

rays  and  speckles 

broad,  reddish 

elongate 

light  pinkish 

on  pinkish  brown 

brown 

mottled 

Number  of  Lots  Examined.  The  total  number  of  lots  and 
the  number  in  each  of  the  following  museum  collections: 
LACM,  AMNH,  ANSP,  MACN,  MNHN,  and  USNM.  These 
counts  also  provide  an  indication  of  the  relative  abundance 
of  each  species. 

Taxonomic  History.  Indicates  whether  authors  have 
understood  the  species  with  the  limits  now  recognized,  or  if 
the  present  treatment  is  a departure  from  past  classifications. 

Abundance  and  Use.  The  fishery  use  and  potential  of  each 
species,  and  the  common  name,  as  provided  by  M.  Bretos. 

Characteristics  and  Variability.  The  first  statement  under 
this  heading  is  a brief  description  meant  to  convey  the  most 
characteristic  features  of  the  species,  followed  by  an  assess- 
ment of  the  variability,  and  whether  variation  may  be  cor- 
related with  geographic  distribution. 

Affinity  and  Comparisons.  As  assessment  of  the  affinity  to 
the  most  closely  related  species  and  the  chief  means  of  dis- 


tinguishing the  species  from  similar  forms.  Remarks  about 
affinity  are  inferences.  Future  work  may  support  or  contradict 
these  suggestions. 

Synonymy  and  Types.  Reasons  for  the  assignment  of  each 
taxon;  the  type  locality,  measurements,  and  repository  of  type 
specimens.  Lectotypes  are  designated  for  lots  consisting  of 
more  than  one  original  specimen. 

SYSTEMATICS 

Family  Fissurellidae  Fleming,  1822 

I recognize  two  subfamilies  in  the  Fissurellidae:  the  Emar- 
ginulinae,  which  I further  subdivide  into  tribes  (McLean, 
1984b,  and  other  work  in  progress),  and  the  Fissurellinae. 

The  Fissurellinae  are  the  youngest  members  of  the  family, 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  17 


Figures  17  and  18.  Fossil  specimens  of  Fissurella  spp.  of  Pliocene  age,  Cerro  Costino,  12  km  S of  Antofagasta,  Chile  (23°45'  S,  70°26'  W), 
collected  by  E,  Martinez.  Specimens  have  lost  the  interior  aragonitic  layer.  (17)  F.  concolor  Philippi,  1887.  LACM  90797 , 60.6  x 40.8  x 9.5 
mm.  (18)  Fissurella  sp.,  cf.  F.  crassa,  LACM  90798,  25.2  x 24.4  x 8.6  mm  (specimen  incomplete). 


appearing  in  the  Cenozoic;  the  Emarginulinae  arose  in  the 
Mesozoic,  with  a burst  of  radiation  in  the  Jurassic. 

Subfamily  Fissurellinae  Fleming,  1822 

DIAGNOSIS.  Apex  of  mature  shell  wholly  absorbed  by 
the  foramen;  foramen  bordered  inside  by  a ring  of  callus  that 
is  not  truncated  or  excavated  posteriorly.  Selenizone  not 
present  in  juvenile  stage.  Shell  muscle  and  muscle  scar  lacking 
intumed  hooked  processes.  Rachidian  tooth  of  radula  narrow 
at  the  tip,  its  base  broad,  no  larger  than  the  adjacent  laterals; 
the  massive  fifth  lateral  usually  with  four  cusps;  marginals 
numerous. 

The  subfamily  Fissurellinae  differs  from  the  subfamily 
Emarginulinae  in  radular  characters  and  in  the  morphology 
of  the  shell  muscle.  In  the  Emarginulinae  the  rachidian  tooth 
is  rhomboidal  in  shape  (whether  broad  or  narrow)  and  the 
enlarged  fifth  lateral  has  only  two  prominent  cusps;  in  the 
Fissurellinae  the  rachidian  tooth  is  narrow  at  the  tip  and  the 
enlarged  fifth  lateral  has  four  cusps  (except  three  in  Ambly- 
chilepas,  one  very  small).  Also,  the  muscle  scar  and  corre- 
sponding shell  muscle  of  the  fissurelline  genera  lack  the  hook- 
shaped process  of  such  emarginuline  genera  as  Puncturel/a 
and  Diodora.  Early  stages  of  emarginuline  genera  have  a 
selenizone  that  may  be  lost  in  mature  stages,  but  the  sele- 
nizone is  not  present  at  any  stage  in  the  Fissurellinae. 

Other  genera  in  the  subfamily  Fissurellinae  are  Ambly- 
chilepas  Pilsbry,  1890,  and  Macrochisma  Sowerby,  1839, 
both  of  which  are  limited  to  the  Indo-Pacific,  and  have  an- 
imals too  large  to  be  contained  in  the  shell. 

Genus  Fissurella  Bruguiere,  1789 

DIAGNOSIS.  Shell  conical,  sculpture  chiefly  radial;  fo- 
ramen nearly  central. 

Two  subgenera  of  Fissurella  are  here  recognized:  Fissu- 
rella, sensu  stricto,  with  a two-layered  shell,  the  outer  layer 
composed  of  calcite,  the  inner  layer  of  aragonite,  and  Cre- 
mides  H.  and  A.  Adams,  1854,  in  which  the  shell  is  composed 


entirely  of  aragonite  (Fig.  30).  This  is  the  first  time  that  the 
subgenera  have  been  so  defined.  As  discussed  above,  it  has 
not  previously  been  noted  that  the  shell  margin  is  indicative 
of  a two-layered  shell  rather  than  merely  a color  difference. 

Recent  authors  (e.g.,  Keen,  in  Knight  et  al.,  1960)  have 


Figure  19.  Distribution  of  Fissurella,  sensu  stricto,  in  North  and 
South  America. 


18  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


regarded  Fissurella,  sensu  stricto,  as  monotypic,  and  have 
placed  the  tropical  species  lacking  the  shell  margin  in  Cre- 
mides  and  the  South  American  species  having  the  shell  mar- 
gin in  Balboaina  Perez-Farfante,  1943.  In  the  present  treat- 
ment, the  scope  of  the  subgenus  Cremides  remains  unchanged, 
but  Balboaina  is  synonymized  with  Fissurella , sensu  stricto. 

Cremides  has  a fossil  record  from  the  Oligocene  (Keen,  in 
Knight  et  al.,  1960),  whereas  Fissurella,  sensu  stricto,  dates 
from  the  Pliocene. 

Subgenus  Fissurella  Bruguiere,  1789 

Fissurella  Bruguiere,  1789:xiv  (genus  without  named  species). 
Type  species,  by  subsequent  monotypy  of  Lamarck,  1799: 
Patella  nimbosa  Linnaeus,  1758. 

Balboaina  Perez-Farfante,  1943:2.  Type  species,  by  original 
designation,  Fissurella  picta  (Gmelin,  1791). 

Carcellesia  Perez-Farfante,  1952:31.  Type  species,  by  orig- 
inal designation,  Fissurella  ( Carcellesia ) doellojuradoi  Per- 
ez-Farfante, 1952  [=  F.  oriens  Sowerby,  1835]. 

Corrina  Christiaens,  1973:93.  Type  species,  by  original  des- 
ignation, Fissurella  ( Corrina ) alba  Philippi,  1845  [=  F. 
oriens  Sowerby,  1835]. 

DIAGNOSIS.  Shell  moderately  large,  composed  of  pig- 
mented, outer  calcitic  layer  that  forms  broad  margin  on  inner 
side;  interior  aragonitic  layer  relatively  thin  except  in  apical 
area,  extending  to  shell  margin  only  in  some  fully  mature 
specimens.  Mantle  papillae  well  developed;  foot  side  tuber- 
cles well  developed,  epipodial  tentacles  short  and  stubby. 

The  species  of  Fissurella,  sensu  stricto,  total  15:  the  13 
species  of  Peru  and  Chile  treated  in  detail  here,  plus  the 
Caribbean  type  species  F.  nimbosa  (Figs.  20-25)  and  the 
Californian  F.  volcano  Reeve,  1 849  (Figs.  26-29).  The  dis- 
tribution of  Fissurella,  sensu  stricto,  is  shown  in  Figure  19. 

Generic  Synonymy.  Bruguiere’s  introduction  of  the  name 
without  reference  to  species  has  caused  some  confusion  over 
the  type  designation.  Although  many  authors  have  followed 
Pilsbry  (1890)  in  the  usage  of  F.  picta  as  the  type  species, 
more  recent  authors  have  followed  Wenz  (1938)  in  citing  F. 
nimbosa,  by  subsequent  monotypy  of  Lamarck  (1799). 

Fissurella  nimbosa  (Figs.  20-25)  is  unlike  all  other  tropical 
species  of  the  genus.  Pilsbry  (1890)  described  the  interior  of 
F.  nimbosa  as  having  a “black  line  around  the  edge.”  He 
also  observed  that  the  species  “has  more  the  aspect  of  the 
Chilean  species  than  that  of  the  West  Indian,”  a comment 
overlooked  by  subsequent  authors.  Fissurella  nimbosa  is  the 
only  species  with  the  two-layered  shell  that  has  a tropical 
distribution.  It  differs  from  all  the  Peruvian  and  Magellanic 
species  in  having  a green  suffusion  to  the  interior,  a character 
shared  with  the  Californian  F.  volcano  (Figs.  26-29). 

Perez-Farfante  (1943)  did  not  notice  that  young  shells  of 
F.  nimbosa  have  a well-defined  dark  margin.  The  shell  she 
figured  (1943,  pi.  1,  figs.  1,  2)  is  mature,  with  the  aragonitic 
layer  obliterating  the  dark  border.  She  considered  Fissurella 
to  be  a monotypic  subgenus  consisting  only  of  F.  nimbosa 
for  reasons  not  clearly  stated,  but  presumably  because  F. 
nimbosa  is  larger  and  more  conical  than  most  of  the  tropical 
species.  She  therefore  missed  the  affinity  of  F.  nimbosa  with 


the  Peruvian  and  Chilean  species  and  proposed  the  subgenus 
Balboaina  for  those  species  with  the  dark  border. 

In  my  opinion,  F.  nimbosa  is  sufficiently  similar  to  other 
species  having  the  two-layered  shell  that  it  can  not  be  sep- 
arated from  them  on  a subgeneric  level.  It  is  premature  to 
offer  a final  opinion  about  the  subgeneric  division  of  the 
group,  and  I am,  therefore,  following  a conservative  course 
in  uniting  those  with  the  calcitic  layer.  Further  evidence  about 
the  affinity  of  the  species  with  the  calcitic  outer  layer  needs 
to  be  offered  from  other  lines  of  investigation,  for  example, 
electrophoresis.  Until  convincing  arguments  can  be  ad- 
vanced to  separate  the  Chilean  species  from  F.  nimbosa  and 
F.  volcano,  I am  treating  Balboaina  as  a synonym  of  Fis- 
surella, sensu  stricto. 

Perez-Farfante’s  original  diagnosis  of  Balboaina  included 
the  following  provisions:  “Margin  of  the  shell  entirely  in  one 
plane,  simple,  not  crenulated  . . . .”  Neither  of  these  traits  is 
true  for  the  majority  of  the  southern  species.  Those  species 
having  radial  ribbing  are  in  fact  finely  crenulate  at  the  margin, 
and  most  are  raised  at  the  sides. 

Two  other  subgeneric  names  have  been  proposed:  Car- 
cellesia Perez-Farfante,  1952,  and  Corrina  Christiaens,  1973. 
Both  names  are  based  on  type  species  that  are  here  regarded 
as  synonyms  of  F.  oriens.  Carcellesia,  type  species  F.  doel- 
lojuradoi Perez-Farfante,  1952,  was  based  upon  a single  aber- 
rant example  of  F.  oriens  with  prominently  raised  ends.  Cor- 
rina. tvDe  species  F.  alba  Philippi,  1845,  was  intended  to 
apply  to  thin-shelled  forms  lacking  sculpture.  I consider  F. 
alba  a synonym  of  F.  oriens,  based  upon  the  scarce  white- 
shelled  form  of  the  species.  Christiaens  also  included  in  Cor- 
rina the  South  African  species  F.  mutabilis  Sowerby,  1835, 
but  that  species  has  no  dark  margin  and  is  therefore  unre- 
lated. Fissurella  oriens  is  a readily  recognizable  species  in 
which  the  sculpture  is  weak,  but  I find  no  reason  to  single  it 
out  as  representing  a subgenus.  Consequently,  both  names 
are  relegated  to  the  synonymy  of  Fissurella,  sensu  stricto. 

Species  Groups.  Pilsbry  (1890)  placed  the  species  having 
a “distinct  dark  marginal  border  inside”  in  four  groups  of 
species  based  on  shell  characters.  I recognize  a somewhat 
similar  scheme  of  three  groups  within  the  subgenus  Fissu- 
rella, sensu  stricto,  based  on  the  presence  or  absence  of  com- 
plex radial  sculpture  and  the  relative  thickness  of  the  calcitic 
and  aragonitic  shell  layers.  Affinity  among  the  species  within 
each  group  is  inferred,  but  this  should  be  tested  by  future 
workers. 

1 . Aragonitic  layer  as  thick  as  calcitic  layer  

Group  of  F.  peruviana 

Aragonitic  layer  markedly  thinner  than  calcitic  layer 
2 

2.  Sculpture  of  primary  and  secondary  ribs  

Group  of  F.  maxima 

Sculpture  of  broad  primary  ribs  only  

Group  of  F.  limbata 

Group  of  Fissurella  peruviana 

Relatively  small-shelled  species  in  which  the  thickness  of  the 
aragonitic  layer  equals  or  approximates  that  of  the  external 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  19 


Figures  20  through  30.  Shells,  radulae,  mantle  lobes,  and  cut  shells  of  Fissurella  nimbosa  (Linnaeus,  1758),  F.  volcano  Reeve,  1849,  and  F. 
( Cremides ) virescens  Sowerby,  1835.  Figures  20  through  25.  F.  nimbosa.  (20)  Cut  shell,  showing  thin,  dark  calcitic  layer.  LACM  76-30,  Puerte 
La  Cruz,  Venezuela,  length  of  cut  edge  20.5  mm.  (21)  Radular  ribbon.  Same  locality,  width  of  ribbon,  1.4  mm.  (22)  Shell,  Frigate  Bay,  St. 
Christopher,  U.S.  Virgin  Islands.  LACM  76-25,  27.0  x 24.9  x 8.1  mm.  (23)  Shell,  Cabo  Blanco,  Isla  Margarita,  Venezuela,  showing  partial 
loss  of  the  external  calcitic  layer  and  wear  obliterating  the  dark  margin  in  beach-worn  shell.  LACM  76-28,  37.9  x 26.2  x 14.0  mm.  (24) 
Mantle  edge.  Frigate  Bay,  St.  Christopher,  U.S.  Virgin  Islands.  LACM  76-25,  length  3.5  mm.  (25)  SEM  view  of  radula.  LACM  76-30,  Puerte 
La  Cruz,  Venezuela,  width  of  field  0.8  mm.  Figures  26  through  29.  Fissurella  volcano.  (26)  Cut  shell,  showing  thin  calcitic  layer.  LACM  66- 
1,  Santo  Tomas,  Baja  California,  Mexico,  length  of  cut  edge  20  mm.  (27)  Mantle  edge.  AHF  1595-47,  Rio  Santo  Tomas,  Baja  California, 


20  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


calcitic  layer;  the  shell  margin  (calcitic  layer)  narrow  at  all 
growth  stages  and  in  mature  specimens  often  obliterated  by 
encroachment  of  the  aragonitic  layer.  Radial  sculpture  of 
primary  and  secondary  ribs,  which  remain  pronounced  at  all 
growth  stages. 

This  group  differs  from  both  the  group  of  F.  maxima  and 
the  group  of  F.  limbata  in  its  relatively  smaller  size  and  in 
having  a thicker  aragonitic  layer  and  a relatively  thin  and 
narrow  calcitic  layer. 

In  addition  to  F.  peruviana,  this  group  includes  the  type 
species  of  Fissurella,  F.  nimbosa  (Linnaeus,  1757),  of  the 
tropical  Caribbean  faunal  province  (Figs.  20-25),  and  F.  vol- 
cano Reeve,  1 849,  of  the  warm  temperate  Californian  Faunal 
Province  (Figs.  26-29).  None  of  the  species  extends  into  the 
cold  temperate  Magellanic  Faunal  Province. 

Although  F.  radiosa  has  an  unusually  narrow  margin,  as 
do  the  members  of  this  group,  it  does  not  have  a sufficiently 
thick  aragonitic  layer  to  suggest  that  it  is  related  to  these 
species. 

In  having  a relatively  thick  aragonitic  layer,  this  group  of 
species  represents  the  connecting  link  between  the  tropical 
species  lacking  the  calcitic  layer  (subgenus  Cremides),  and 
the  South  American  species  with  thick  calcitic  layers  (sub- 
genus Fissurella,  sensu  stricto). 

Fissurella  peruviana  Lamarck,  1822 

Figures  31-50 

Fissurella  peruviana  Lamarck,  1 822,  6(2):  1 5;  Orbigny,  1841: 
74;  Delessert,  1841,  pi.  24,  fig.  7;  Reeve,  1849,  pi.  5,  figs. 
26a-d;  Hupe,  1854:241;  Philippi,  1860:181;  Sowerby  II, 
1862:185,  figs.  38-41;  Pilsbry,  1890:155,  pi.  33,  figs.  41- 
45,  pi.  42,  figs.  57-59;  Dali,  i 909: 178,  242;  Mermod,  1 950: 
713,  fig.  22;  Riveros-Zuniga,  195 1 : 1 30,  fig.  35;  Pena,  1970: 
156;  Dell,  1971:190;  Christiaens,  1973:86;  Ramirez- 
Boehme,  1974:31  [key], 

Fissurella  subrotunda  Deshayes,  1830:135;  Deshayes  in  La- 
marck, 1836,  7:602;  Orbigny,  1841:74  [under  F.  peru- 
viana];  Reeve,  1849,  pi.  5 [under  F.  peruviana}. 

Fissurella  affinis  “Gray,”  Sowerby,  1835a:  125;  Sowerby, 
1835b:4,  fig.  44;  Sowerby  II,  1862:185,  figs.  46,  179;  Chris- 
tiaens, 1973:83. 

Fissurella  clypeus  Sowerby,  1835a:  128;  Sowerby,  1835b:4, 
fig.  44;  Reeve,  1850,  fig.  76;  Sowerby  II,  1862:185,  fig.  63; 
Pilsbry,  1890:156,  pi.  60,  fig.  82,  pi.  31,  fig.  20;  McLean 
in  Keen,  1971:901;  Christiaens,  1973:83  [under  F.  asperel- 
la], 

Fissurella  occidens  Gould,  1846:156;  Gould,  1852:364,  pi. 
31,  figs.  473a,  b;  Pilsbry,  1890:155  [under  F.  peruviana]-, 
Johnson,  1964:118  [holotype,  USNM  5863], 

Fissurella papudana  Ramirez-Boehme,  1 974: 1 8,  3 1 [key],  pi. 
2,  figs.  5a,  b,  c. 


SheSL  Relatively  small  (25-40  mm  mature  length);  vari- 
able in  height  from  low  to  high  conical;  variable  in  outline 
from  broadly  oval  to  elongate  and  uneven;  plane  of  margin 
also  varying  and  probably  conforming  to  an  attachment  site. 
Sculpture  of  fine,  often  imbricate,  radial  ribs,  primary  ribs 
remaining  strong.  Color  highly  variable,  including  some  that 
are  solid  dark  red  to  reddish  gray,  some  that  are  faintly  rayed, 
and  some  with  rays  of  brown  and  white;  lateral  rays  of  elon- 
gate specimens  often  curving  forward.  Margin  relatively  nar- 
row at  all  growth  stages,  reflecting  pattern  of  rays  through 
full  thickness  of  calcitic  layer.  Cut  shells  show  aragonitic  layer 
slightly  thicker  than  calcitic  layer.  Foramen  elongate  and 
tripartite  in  earliest  stages  but  quickly  becoming  oval  in  ju- 
venile shells  and  broadly  oval  in  mature  shells. 

Juvenile  Shell.  Strongly  sculptured,  conical;  frequently 
reddish  with  two  lateral  white  rays.  Mature  specimens  with 
brown  and  white  rays  have  a red  ring  in  the  calcitic  layer 
surrounding  the  foramen,  a remnant  of  the  juvenile  red  phase. 

Mantle  and  Foot.  Fully  retractable  in  the  shell.  Mantle 
banded  to  match  the  rays  of  the  shell.  Mantle  lobe  relatively 
narrow,  upper  and  lower  edges  with  branched  papillae.  Foot 
side  brown  to  black,  with  numerous,  projecting,  light-tipped 
tubercles. 

Habitat.  Lower  intertidal  zone  to  20  m,  but  most  abundant 
in  the  sublittoral  zone.  Scattered  individuals  occurring  at  low 
tide,  nestled  in  crevices  on  the  sides  of  rocks,  the  shell  outline 
moulded  to  fit  the  site  of  attachment.  Occurring  at  greater 
depths  than  any  other  of  the  species  in  northern  Chile.  At 
Mejillones  (23°02'  S)  it  was  the  only  species  that  I saw  at  10- 
20  m on  a sloping  rocky  bottom  dominated  by  the  mussel 
Aulacomya  ater. 

Distribution.  Chiclayo,  Peru  (6°47'  S)  (LACM,  collector 
unknown;  also  reported  at  Chiclayo  by  Pena,  1970),  to  Gua- 
bun,  northwestern  tip  of  Isla  de  Chiloe,  Chile  (41°50'  S) 
(LACM  75-40,  McLean).  I have  found  specimens  at  most 
localities  between  Isla  Guanape,  Peru  (8°32'  S),  and  Rio  Bio- 
bio, Concepcion  Province,  Chile  (36°48'  S),  but  did  not  find 
it  at  Mehuin,  Valdivia  Province. 

Number  of  Lots  Examined.  1 18  (LACM  52,  AMNH  11, 
ANSP  11,  MACN  3,  MNHN  17,  USNM  24). 

Taxonomic  History.  Fissurella  peruviana  has  been  reason- 
ably well  understood  by  authors,  at  least  with  reference  to 
its  occurrence  in  Peru.  Its  presence  in  Chile  has  not  been 
adequately  discussed;  Riveros-Zuniga  (1951)  merely  listed 
previous  authors  who  cited  records  from  Chile.  A low  form 
of  this  species  with  irregular  outline  is  common  in  Chile  (Figs. 
36,  37,  39-42)  and  was  illustrated  by  Riveros-Zuniga  (1951), 
rnisidentified  as  F.  stellata  Reeve,  1850.  This  form  was  twice 
named  by  Sowerby  in  1835  and  again  by  Ramirez-Boehme 
in  1974.  Christiaens  (1973)  considered  F.  affinis  Sowerby  to 
be  a good  species,  but  there  is  little  justification  for  such  a 


Mexico,  length  5 mm.  (28)  Shell.  White’s  Point,  Los  Angeles  County,  California.  LACM  69-37,  28.7  x 20.0  x 11.7  mm.  (29)  Shell,  same 
locality,  27.2  x 21.0  x 8.3  mm.  (30)  Fissurella  ( Cremides ) virescens,  cut  shell,  calcitic  layer  lacking,  La  Cruz  de  Juanacaxtl,  Nayarit,  Mexico. 
LACM  71-33,  length  of  cut  edge  23.7  mm. 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  21 


Figures  31  through  42.  Fissurella  peruviana  Lamarck,  1822.  Mature  shells.  (31)  3-5  m,  Isla  Guanape,  Peru.  LACM  74-3,  45.1  x 23.5  x 
12.8  mm.  (32)  Holotype,  F.  occidens  Gould.  Callao,  Peru.  USNM  5863,  35  x 27  x 19  mm.  (33)  Lectotype,  F.  subrotunda  Deshayes.  “Peru.” 
MNHNP,  31.3  x 27.4  x 16.8  mm.  (34)  Paracas,  lea  Province,  Peru.  LACM  72-79,  31.5  x 24.0  x 17.8  (beach  shell).  (35)  Laguna  Grande, 
lea  Province,  Peru.  LACM  72-77,  31.2  x 25.9  x 11.5  mm  (beach  shell).  (36)  Iquique,  Chile.  LACM  64-16,  24.0  x 16.7  x 7.6  mm.  (37) 


22  Contributions  in  Science,  Number  354 


MfcLean:  Peruvian  and  Magellanic  Fissurella 


view,  as  discussed  below.  The  broad  distribution  and  the 
extent  of  intraspecific  variation  in  F.  peruviana  have  not 
previously  been  understood. 

Abundance  and  Use.  Because  of  its  chiefly  sublittoral  hab- 
itat, F.  peruviana  seems  to  be  uncommon,  although  beach- 
worn  shells  are  common  throughout  the  range.  This  is  the 
smallest  species  of  Fissureila  in  Peru  and  Chile  and  therefore 
has  little  importance  as  a food  resource.  According  to  M. 
Bretos,  the  species  has  no  common  name  in  northern  Chile, 
perhaps  because  it  is  considered  the  juvenile  form  of  such 
species  as  F.  maxima. 

Characteristics  and  Variability.  Fissureila  peruviana  is 
characterized  by  its  small  size,  imbricate  radial  ribs,  oval 
foramen,  narrow  margin,  and  relatively  thick  aragonitic  lay- 
er. In  lateral  profile  it  varies  from  low  to  extremely  conical; 
in  outline  it  varies  from  broadly  oval  to  elongate  and  irreg- 
ular. Its  color  varies  from  uniformly  dark  reddish  to  gray  or 
rayed  with  brown  and  white.  It  is  the  most  variable  species 
of  Fissureila  in  the  Peruvian  Faunal  Province. 

Specimens  from  Peru  tend  to  be  more  conical  and  more 
uniformly  colored,  whereas  those  from  central  Chile  tend  to 
be  flatter  and  are  more  likely  to  be  rayed  with  brown  and 
white.  However,  I am  not  convinced  that  a geographic  dis- 
tinction can  be  drawn.  I have  noticed  that  the  more  conical 
forms  occur  on  rocks  adjacent  to  sandy  bottoms,  whereas  the 
flattened  forms  with  irregular  outlines  are  found  on  rocky 
bottoms  away  from  sand.  At  Iquique  and  Antofagasta  the 
flattened  irregular  forms  occur,  but  I have  found  some  highly 
conical  forms  associated  with  the  scallop  beds  north  of  An- 
tofagasta at  Bahia  Morena.  Conical  specimens  have  also  been 
seen  from  the  Concepcion  vicinity.  The  recognition  of  geo- 
graphic subspecies  is  therefore  not  justified. 

Affinity  and  Comparisons.  Fissureila  peruviana  differs  from 
all  other  Peruvian  and  Magellanic  species  in  having  a rela- 
tively thick  aragonitic  layer  and  in  having  the  oval  foramen 
well  developed  at  an  early  stage.  Yet  these  differences  do  not 
seem  sufficient  to  regard  it  as  unrelated  to  those  species  with 
well-defined  primary  and  secondary  ribs  in  the  group  of  Fis- 
surella  maxima.  Of  those  species,  it  has  the  most  in  common 
with  F.  costata,  with  which  it  shares  overall  shape  and  the 
small,  oval  foramen.  It  more  closely  resembles  F.  volcano 
from  California,  which  is  also  relatively  small,  with  a narrow 
margin  and  proportionately  thick  aragonitic  layer.  Fissureila 
peruviana  differs  from  F.  volcano  in  having  a more  oval 
foramen  and  in  lacking  the  green  tinge  of  the  interior  and 
pink-bordered  callus  of  F.  volcano.  F.  volcano  is  an  intertid- 
ally  occurring  species  and  F.  peruviana  is  characteristically 
sublittoral. 

When  compared  to  young  specimens  of  other  Peruvian 
and  Chilean  species,  F.  peruviana  can  be  distinguished  by  its 
narrow  margin  and  oval  foramen.  Small  specimens  may  be 


distinguished  from  the  juveniles  of  F.  maxima  in  having  the 
foramen  more  oval  and  the  margin  not  rounded.  The  primary 
ribs  are  not  as  strongly  developed  as  those  in  young  stages 
of  F.  costata,  F.  picta  lata,  or  F.  radiosa. 

Synonymy  and  Types.  Fissureila  peruviana  Lamarck,  1822, 
is  one  of  the  three  earliest-named  species  of  the  region.  La- 
marck’s specimens  were  from  the  “coasts  of  Peru,”  and  some 
of  the  specimens  were  said  by  Lamarck  to  be  less  conical 
than  others.  Unfortunately,  the  first  illustration  of  Lamarck’s 
species  (Delessert,  1841)  may  be  a specimen  of  some  other 
species,  as  suggested  by  Pilsbry  ( 1 890),  Mermod  ( 1 950),  and 
Christiaens  (1973).  Mermod  (1950)  discussed  two  Lamarck- 
ian specimens  in  the  Geneva  Museum  collection  considered 
to  be  original.  “Specimen  no.  2”  discussed  by  Mermod  fits 
the  present  concept  of  F.  peruviana  and  is  inscribed  by  the 
hand  of  Lamarck,  according  to  Mermod.  I am  not  able  to 
determine  the  identity  of  “specimen  no.  1,”  which  is  appar- 
ently the  specimen  figured  by  Delessert.  Inasmuch  as  La- 
marck indicated  that  there  were  several  specimens,  I hereby 
designate  “specimen  no.  2”  as  the  lectotype,  which  is  in 
accord  with  the  original  intent  of  the  author.  The  lectotype 
(see  Mermod,  1950,  fig.  22-2)  represents  the  high  conical 
form  of  F.  peruviana  with  a nearly  circular  foramen;  length 
20,  width  25,  height  16  mm. 

I have  examined  8 syntypes  and  the  original  mounting 
board  of  F.  subrotunda  Deshayes,  1830,  received  on  loan 
from  the  Paris  Museum.  Lengths  are  35.6,  31.3,  30.3,  30.0, 
25.7,  22.7,  and  20.7  mm.  The  original  measurements  were 
32  mm  in  length  and  28  mm  in  width;  the  second  largest 
specimen  is  here  designated  the  lectotype  (Fig.  33);  it  is  31.3 
mm  in  length  and  27.4  mm  in  width  and  is  probably  the 
measured  specimen.  Type  material,  from  “Perou,”  appar- 
ently has  not  previously  been  illustrated.  All  specimens  are 
dark  reddish  rayed,  moderately  conical,  and  with  an  oval 
basal  outline.  No  comparisons  were  originally  made  with  F. 
peruviana;  Orbigny  (1841)  relegated  the  name  to  the  syn- 
onymy of  F.  peruviana. 

Type  material  of  F.  affinis  Sowerby,  1835,  has  not  been 
located.  Reeve  (1849)  regarded  it  as  an  elongate,  flattened 
form  of  F.  peruviana.  Sowerby  II  (1862)  doubtfully  recog- 
nized it  as  a species  “in  order  to  avoid  the  extreme  incon- 
venience of  including  opposite  characters  under  the  same 
name  . . . .”  Christiaens  (1973)  maintained  it  as  a species, 
but  in  my  opinion  the  original  figure  in  the  “Conchological 
Illustrations”  represents  the  extreme  flattened  form  of  F. 
peruviana,  which  occurs  throughout  the  range  of  the  species 
in  Chile.  Several  localities  were  mentioned  originally:  “In- 
sulas Mexillones  et  Lobos,  Iquiqui,  and  Valparaiso.” 

The  holotype  of  F.  clypeus  Sowerby,  1835  (Fig.  40),  was 
said  to  have  come  from  “Sanctam  Elenam,”  presumably  the 
Santa  Elena  Peninsula,  Ecuador.  This  name  has  baffled  sub- 


Iquique,  Chile.  LACM  75-12,  23.6  x 16.4  x 7.4  mm  (beach  shell).  (38)  2-4  m,  El  Rincon  de  Mejillones,  Antofagasta  Province,  Chile.  LACM 
75-23,  32.0  x 29.8  x 15.0  mm.  (39)  2-5  m,  Antofagasta,  Chile.  LACM  75-20,  27.0  x 18.8  x 17.6  mm.  (40)  Holotype,  F.  clypeus  Sowerby. 
Locality  doubtful.  BMNH  197578,  28.0  x 18.9  x 5.8  mm.  (41)  Holotype,  F.  papudana  Ramirez-Boehme.  Papudo,  Aconcagua  Province, 
Chile.  MNHN  200374,  36  x 26  x 10  mm.  (42)  Guabun,  Chiloe  Province,  Chile.  LACM  75-40,  27.3  x 19.7  x 7.9  mm  (beach  shell). 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissureila  23 


Figures  43  through  50.  Fissure/Ia  peruviana  Lamarck,  1822.  Radula,  mantle  lobe,  cut  shell,  juvenile  shells,  and  intact  specimens.  (43)  Radula, 
light  microscope  preparation.  Montemar,  Valparaiso  Province,  Chile.  LACM  75-30,  width  of  ribbon  0.4  mm.  (44)  Radula,  SEM.  7-20  m, 
Mejillones,  Chile.  LACM  75-21,  width  of  field  1 mm.  (45)  Mantle  edge.  1-2  m,  Playa  Hermosa,  Ancon,  Peru.  LACM  74-21,  length  4 mm. 
(46)  Cut  shell.  Los  Colorados,  Antofagasto,  Chile.  LACM  75-19,  length  21  mm.  (47)  Juvenile  shell.  Same  locality.  4.5  x 2.7  x 1.5  mm.  (48) 
Juvenile  shell.  3-5  m,  Isla  Guanape,  Peru.  LACM  74-3,  9.0  x 5.3  x 2.4  mm.  (49)  Body  of  preserved  specimen.  Antofagasta,  Chile.  LACM 
75-15,  shell  length  25.1  mm.  (50)  Living  specimen.  Same  locality,  same  specimen. 


sequent  authors,  especially  because  a view  of  the  internal 
margin  has  never  been  given  and  the  foramen  has  been  fig- 
ured to  be  much  longer  that  it  actually  is.  In  my  opinion  it 
is  the  flattened  form  of  F.  peruviana,  as  was  suspected  by 
Sowerby  II  ( 1 862),  and  the  shell  more  likely  came  from  Chile. 
The  Ecuadorian  locality  is  well  to  the  north  of  the  northern 
limit  of  F.  peruviana,  and  the  flattened  form  of  the  species 
is  particularly  common  in  Chile.  The  foramen  of  the  speci- 
men is  slightly  longer  than  normal.  Pilsbry  (1890)  copied  the 
Reeve  figure,  but  his  figure  82,  said  to  be  a copy  of  the 
Sowerby  figure,  is  not  that,  which  has  contributed  to  the 
confusion. 

The  holotype  of  F.  occidens  Gould,  1846,  is  USNM  5863, 
length  35  mm  (Fig.  32).  There  are  two  paratypes,  MCZ  1 55766 
(Johnson,  1964).  The  type  locality  is  Callao,  Peru.  The  ho- 
lotype represents  the  high-conical,  reddish  rayed  form  so 
abundant  in  Peru. 

Fissurella  papudana  Ramirez-Boehme,  1974,  was  de- 
scribed without  comparisons  to  other  species.  The  type  lo- 


cality was  Papudo,  Aconcagua  province,  Chile.  The  holotype 
(Fig.  41),  MNHN  200374,  is  an  example  of  the  flattened 
form  of  F.  peruviana,  with  a narrow  margin,  irregular  outline, 
oval  foramen,  and  brown  and  white  rays. 

Group  of  Fissurella  maxima 

Relatively  large-shelled  species  in  which  the  thickness  of  the 
exterior  calcitic  layer  of  the  shell  greatly  exceeds  that  of  the 
interior  aragonitic  layer.  All  species  have  strong  radial  sculp- 
ture, at  least  in  the  early  stages.  Sculpture  consists  of  primary 
ribs  and  weaker  secondary  ribs  that  arise  between  the  primary 
ribs.  Mature  shells  may  retain  the  distinction  between  pri- 
mary and  secondary  ribs,  or  all  the  ribs  may  attain  a similar 
strength,  whether  coarse  or  very  fine. 

There  are  eight  species  in  the  group  of  F.  maxima,  four 
in  the  Peruvian  Provice  and  four  in  the  Magellanic  Province. 
Most  of  the  species  in  this  group  are  highly  variable  and 
eurytopic,  tolerant  of  a broad  range  of  conditions  of  intertidal 


24  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


Figures  51  through  55.  Fissurella  maxima  Sowerby,  1835.  Mature  shells.  (51)  Bahia  Independencia,  Peru.  AHF  380-35,  84.0  x 47.7  x 23.0 
mm.  (52)  Lectotype,  F.  hondurasensis  Reeve.  Locality  unknown.  BMNH  1976139,  56.6  x 32.5  x 13.5  mm.  (53)  Iquique,  Chile.  LACM  64- 
16,  26.0  x 14.4  x 5.8  mm.  (54)  Los  Molles,  Aconcagua  Province,  Chile.  LACM  75-28,  73.9  x 47.4  x 18.0  mm.  (55)  Holotype,  F.  maxima 
Sowerby.  Valparaiso,  Chile.  BMNH  197569,  128.8  x 85.4  x 34.7  mm. 


exposure.  Most  species,  particularly  those  with  high  vari- 
ability, have  been  ovemamed,  the  synonyms  applying  to 
variant  specimens. 

Only  one  species  in  this  group,  F.  nigra,  is  stenotopic,  with 
a narrow  ecological  tolerance,  and  low  variability.  It  also 
differs  from  the  other  in  having  primary  and  secondary  ribs 
pronounced  only  in  juvenile  stages. 


Three  of  the  four  Magellanic  species  in  this  group  have 
geographic  subspecies. 

Fissurella  maxima  Sowerby,  1835 

Figures  51-63 

Fissurella  maxima  Sowerby,  1835a:  123;  Sowerby,  1 8 3 5b: 3, 
fig.  18;  Orbigny,  1841:475,  pi.  64,  figs.  4-7;  Philippi,  1845: 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  25 


Figures  56  through  63.  Fissurella  maxima  Sowerby,  1835.  Radula,  mantle  lobe,  cut  shell,  juvenile  shell,  and  living  specimens.  (56)  Mantle 
lobe.  Iquique,  Chile.  LACM  75-12,  length  9 mm.  (57)  Radular  teeth,  lateral  view  of  large  outer  lateral  teeth.  Iquique,  Chile.  LACM  70-68, 
width  of  ribbon  0.4  mm,  shell  length  20.5  mm.  (58)  Radular  ribbon,  same  specimen.  Width  of  field  1.0  mm.  (59)  Living  specimen,  anterior 
end,  showing  head  with  cephalic  tentacles  and  foot  side.  Iquique,  Chile.  LACM  75-12.  (60)  Juvenile  specimen.  Punta  El  Lacho,  Santiago 
Province,  Chile.  LACM  75-32,  14.0  x 7.8  x 2.9  mm.  (61)  Radular  ribbon,  air-dried.  Bahia  San  Juan,  Peru.  AHF  828-38,  width  of  ribbon 
2.5  mm,  shell  length  58.9  mm.  (62)  Living  specimen  on  rock  substrate,  anterior  at  right.  Pozo  Toyo,  Tarapaca  Province,  Chile.  LACM  75- 
10.  (63)  Cut  shell.  Montemar,  Valparaiso  Province,  Chile.  LACM  75-30,  length  of  cut  edge  37  mm. 


3,  pi.  1,  fig.  1;  Reeve,  1849,  pi.  4,  fig.  22;  Hupe,  1854:239; 
Philippi,  1860:180;  Sowerby  II,  1862:187,  figs.  8,  9;  Wat- 
son, 1886:33;  Pilsbry,  1890:145,  pi.  30,  figs.  8,  9,  pi.  33, 
figs.  46,  47;  Dali,  1909:242;  Ziegenhom  and  Thiem,  1925: 
11,  pi.  1,  figs.  8a,  8b,  9a,  9b;  Pena,  1970:156;  Riveros- 
Zufiiga,  1951:102,  fig.  18;  Dell,  1971:188,  pi.  4,  figs.  1-3; 
Marincovich,  1973:18,  fig.  31;  Ramirez-Boehme,  1974:31 
[key]. 

Fissurella  solida  Philippi,  1845:142;  Carcelles  and  William- 
son, 1951:  256;  Dell,  1971:193;  Ramirez-Boehme,  1974: 
31  [key]. 


Fissurella  concinna  Philippi,  1845:143;  Philippi,  1846:66, 
pi.  2,  fig.  5;  Reeve,  1850,  pi.  15,  fig.  112;  Hupe,  1854:245; 
Sowerby  II,  1862:187,  figs.  4,  178,  206;  Rochebrune  and 
Mabille,  1889:71;  Pilsbry,  1890:146,  pi.  32,  figs.  32,  33, 
pi.  45,  figs.  7,  8 [as  var.  of  F.  maxima ];  Carcelles  and 
Williamson,  1951:256;  Pena,  1970:156;  Dell,  1971:183; 
Ramirez-Boehme,  1974:30  [key]. 

Fissurella  hondurasensis  Reeve,  1849,  pi.  7,  fig.  48;  Pilsbry, 
1890:146,  pi.  35,  fig.  6. 

Shell.  Relatively  large  (80  to  135  mm  mature  length),  low 


26  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


to  medium  in  height,  outline  elongate-oval;  sides  of  shell 
elevated.  Radial  ribs  prominent,  low  and  rounded,  alternat- 
ing in  strength,  crenulating  margin.  Ground  color  pale  yellow; 
rays  dark  purple,  broad,  uninterrupted;  primary  ribs  centered 
in  each  light  and  dark  ray.  Margin  very  broad  in  growing 
shells,  narrow  in  mature  shells,  rounded  at  junction  with 
aragonitic  layer;  margin  zoned;  outer  zone  narrow,  showing 
color  rays;  inner  zone  broader,  uniformly  pale  and  translu- 
cent. Cut  shells  show  pigment  of  rays  concentrated  at  surface. 
Foramen  nearly  central,  elongate  in  young  shells,  oval  in 
mature  shells. 

Juvenile  Shell.  High  conical,  base  elongate,  margin  round- 
ed; ribs  rounded,  strong;  color  reddish,  with  two  lateral  white 
rays  and  scattered  black  flecks.  After  length  of  6 mm  shell 
becomes  flatter,  the  primary  ribs  nodulous  and  much  more 
prominent  than  the  secondary  ribs;  the  light  and  dark  rayed 
pattern  emerges  and  the  black  flecks  are  lost. 

Mantle  and  Foot.  Not  fully  retractable  in  shell;  cephalic 
tentacles  brown,  tipped  with  yellow.  Mantle  lobe  broad, 
banded  with  purplish-brown  and  light  gray  to  match  rays  on 
shell;  papillae  of  upper  edge  bulbous,  finely  branched,  those 
of  lower  edge  smaller.  Side  of  foot  brown;  tubercles  strongly 
developed,  tips  lighter  colored. 

Habitat.  Partially  exposed  rocky  areas  at  low  tide  to  4 m. 
Some  individuals  are  free  of  encrusting  algae  and  live  on  the 
under  sides  of  large  flat  rocks  at  low  tide,  but  most  live  in 
the  open  and  are  covered  with  a dense  low  growth  of  red 
algae.  When  exposed  at  low  tide  they  are  tightly  wedged  in 
crevices.  Only  rarely  do  specimens  have  attached  Scurria 
parasitica  on  the  shell. 

Distribution.  Huarmey,  Peru  (10°06'  S)  (LACM  70-97,  E 
del  Solar),  to  Lirquen,  Concepcion  Province,  Chile  (36°41' 
S)  (LACM  72-207,  Univ.  Concepcion).  The  southermost  range 
of  F.  maxima  in  Chile  is  not  as  extensive  as  that  of  most  of 
the  other  common  species  of  the  Peruvian  Faunal  Province. 
I found  no  trace  of  it  at  Rio  Bio-bio,  Concepcion  Province, 
Mehuin,  near  Valdivia,  or  Guabun  on  the  northwest  tip  of 
Isla  de  Chiloe.  More  extreme  records  in  the  literature  are 
rejected:  Dali’s  (1909)  record  from  Manta,  Ecuador,  and  Ri- 
veros-Zuniga’s  (1951)  record  from  Fuerte  Bulnes  in  the  Strait 
of  Magellan.  Dell’s  (1971)  records  from  Isla  de  Chiloe  are 
based  upon  specimens  of  F.  picta  lata. 

Number  of  Lots  Examined.  128  (LACM  34,  AMNH  25, 
ANSP  7,  MACN  14,  MNHN  30,  USNM  18). 

Taxonomic  History.  Fissurella  maxima  has  been  recog- 
nized by  previous  authors.  Its  three  synonyms  have  not  been 
considered  important,  although  F.  concinna  Philippi  has 
sometimes  been  given  status  as  a narrow  “form”  of  the  species. 

Abundance  and  Use.  Common  throughout  its  range  and 
particularly  abundant  in  central  and  northern  Chile.  It  is  one 
of  the  major  species  used  for  food,  taken  by  shore  collectors 
and  divers  in  relatively  shallow  water.  It  is  called  the  “lapa 
de  huiros,”  because  it  frequently  occurs  near  the  brown  algae 
Lessonia,  known  as  “huiros.” 

Characteristics  and  Variability.  Always  strongly  ribbed  and 
having  a consistent  color  pattern  of  dark  purple  rays,  the 
most  characteristic  feature  is  the  interior  border,  which  is 


uniquely  rounded  and  has  two  zones.  The  margin  is  always 
crenulated  by  the  radial  ribs,  even  in  the  largest  specimens. 
It  is  not  a highly  variable  species;  color  pattern  and  shell 
height  are  consistent.  Width  of  the  shell  is  the  most  variable 
feature;  some  specimens  may  be  especially  elongate  (Fig.  5 1 ); 
oval  specimens  are  rare.  Largest  specimens  come  from  shell 
piles  in  central  Chile;  those  from  Peru  are  small.  Elongate 
specimens  are  frequently  those  from  Peru.  However,  I do 
not  consider  the  geographic  differences  of  sufficient  impor- 
tance to  warrant  recognition  of  subspecies. 

Affinity  and  Comparisons.  Fissurella  maxima  has  features 
that  represent  an  extreme;  its  rounded  margin  is  not  shared 
by  other  species.  It  most  resembles  F.  cumingi,  which  has  a 
similarly  large  mantle  and  foot,  and  similar  size,  height,  and 
color  pattern.  Fissurella  maxima  differs  in  having  a rounded 
margin,  stronger  sculpture,  and  more  pronounced  primary 
ribbing.  It  might  also  be  confused  with  F.  picta  lata,  but  it 
is  lower,  more  elongate,  and  has  broader  ribs  than  that  species. 
Juveniles  are  reddish  like  those  of  F.  peruviana,  but  differ  in 
having  an  upturned  margin,  a more  elongate  foramen,  and 
are  more  elongate. 

Synonymy  and  Types.  Fissurella  maxima  was  described 
by  Sowerby,  1835,  from  a specimen  collected  by  Cuming  at 
Valparaiso,  Chile.  The  holotype,  BMNH  197569  (Fig.  55), 
length  128.8  mm,  matches  the  specimen  figured  by  Sowerby 
(1835b).  Reeve  (1849)  figured  a different  specimen;  these  two 
specimens  have  also  been  figured  by  Dell  (1971,  pi.  4,  fig.  1, 
holotype;  figs.  2,  3,  Reeve  specimen). 

I have  not  located  any  type  material  of  F.  solida  Philippi, 
1845,  from  “Chile.”  It  has  not  been  illustrated.  Philippi’s 
description  mentions  what  I interpret  as  the  rounded  internal 
border  of  F.  maxima  and  states  that  he  did  not  know  the 
young  of  F.  maxima;  other  features  are  in  accord  with  F. 
maxima,  so  I am  confident  that  the  name  should  be  relegated 
to  the  synonymy  of  F.  maxima. 

Type  material  of  F.  concinna  Philippi,  1845,  also  from 
“Chile,”  has  not  been  located.  Philippi’s  figures  have  sug- 
gested to  most  authors  that  it  is  a small,  laterally  compressed 
form  of  F.  maxima.  As  with  F.  solida,  Philippi  noted  the 
rounded  margin.  Stunted,  narrow  specimens  matching  his 
figure  are  present  in  collections;  those  from  central  Peru  (Fig. 
51)  may  be  predominantly  narrow,  but  broader  specimens 
occur  at  the  same  localities  and  it  is  doubtful  that  the  name 
has  any  taxonomic  utility.  Rochebrune  and  Mabille  (1889) 
used  the  name  incorrectly  for  specimens  from  Tierra  del 
Fuego  (no  doubt  confusing  it  with  F.  radiosa );  this  accounts 
for  the  Fuegan  records  of  the  “variety”  concinna  repeated 
by  subsequent  authors. 

Fissurella  hondurasensis  Reeve,  1 849,  supposedly  (in  error) 
from  “Honduras,”  was  correctly  placed  in  the  synonymy  of 
F.  maxima  by  Pilsbry  (1890).  There  are  four  specimens  with 
the  original  mounting  board  now  labeled  syntypes,  BMNH 
1976139,  lengths  56.6,  36.8,  33.0,  and  28.5  mm;  the  largest 
is  here  figured  (Fig.  52)  and  designated  the  lectotype.  All  are 
young  specimens,  more  thin-shelled  and  with  narrower  mar- 
gins than  usual,  but  the  dark  flecks  of  the  juveniles  of  F. 
maxima  are  apparent,  and  I am  confident  of  their  identity 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  27 


with  F.  maxima.  A fifth  specimen  originally  mounted  on  the 
same  board  is  a young  specimen  of  F.  pulchra. 

Fissure! la  latimarginata  Sowerby,  1835 

Figures  64-79 

Fissurella  latimarginata  Sowerby,  1835a:  126;  Sowerby, 
1835b:3,  fig.  69;  Gray,  1839: 148,  pi.  39,  fig.  8;  Reeve,  1849, 
pi.  3,  fig.  1 9;  Hupe,  1 854:242;  Philippi,  1 860: 1 80;  Sowerby 
II,  1862:185,  figs.  6,  7,  12;  Pilsbry,  1890:153,  pi.  32,  figs. 
36-38;  Dali,  1909:242;  Ziegenhom  and  Thiem,  1925:17, 
pi.  2,  figs.  22a,  22b;  Carcelles  and  Williamson,  1951:255; 
Riveros-Zuniga,  1951:125,  fig.  33;  Pena,  1970:156;  Dell, 
1971:187,  pi.  3,  figs.  12-14;  Marincovich,  1973:17,  fig.  29; 
Ramirez-Boehme,  1974:31  [key]. 

Fissurella  biradiata  Sowerby,  1 835a:  1 24;  Sowerby,  1 835b;3, 
figs.  23,  52;  Orbigny,  1841:477;  Reeve,  1849,  pi.  3,  fig.  20; 
Philippi,  1860:180;  Sowerby  II,  1862:185,  figs.  1-3;  Zie- 
genhom and  Thiem,  1925:17,  pi.  2,  fig.  23;  Dell,  1971: 
182,  pi.  3,  figs.  10,  11. 

Fissurella  latimarginata  var.  biradiata,  Pilsbry,  1 890: 1 54,  pi. 
35,  fig.  3,  pi.  46,  figs.  12-14;  Riveros-Zuniga,  1951:126, 
fig.  34;  Ramirez-Boehme,  1974:31  [key], 

Fissurella  bella  Reeve,  1849,  pi.  3,  fig.  21;  Sowerby  II,  1862: 
185,  fig.  25;  Pilsbry,  1890:150,  pi.  33,  fig.  48;  Riveros- 
Zuniga,  1951:119;  Dell,  197 1 : 182,  pi.  3,  figs.  3, 4;  Ramirez- 
Boehme,  1974:32  [key], 

Fissurella  galericulum  Reeve,  1850,  pi.  11,  fig.  77. 
Fissurella  latimarginata  var.  galericulum,  Ramirez-Boehme, 
1974:31  [key], 

Fissurella  punctatissima  Pilsbry,  1890:150,  pi.  58,  figs.  21- 
23;  Dali,  1909:124;  Riveros-Zuniga,  1951:118,  fig.  26;  Dell, 
1971:192. 

Shell.  Relatively  large  (70  to  1 15  mm  mature  length);  low 
to  moderately  high;  outline  oval,  markedly  tapered  ante- 
riorly; base  of  shell  in  one  plane  or  slightly  elevated  on  sides. 
Sculpture  of  very  fine,  sharply  raised  radial  ribs.  Color  uni- 
formly dark  purplish  red  to  gray,  except  for  two  lighter  rays 
extending  laterally  in  young  shells,  fading  away  in  mature 
shells;  occasional  specimens  faintly  rayed  throughout.  Mar- 
gin very  broad  and  flat  in  young  shells,  solid  dark  red,  outer 
edge  with  narrow  gray  zone.  Cut  shells  showing  gray  outer 
zone  in  calcitic  layer;  recent  growth  of  outermost  zone  in 
some  mature  shells  changing  from  gray  to  nearly  colorless. 
Foramen  very  long  and  tripartite  in  young  shells,  elongate- 
oval  in  mature  shells. 

Juvenile  Shell.  Elongate  and  elevated,  radial  ribs  faint, 
nearly  black  except  for  two  lateral  white  rays.  After  shell 


length  of  5 mm,  new  growth  less  conical,  fine  radial  ribs 
stronger,  and  shell  lighter  in  color,  not  yet  showing  gray  outer 
zone  to  calcitic  layer. 

Mantle  and  Foot.  Not  fully  retractable  in  shell;  mantle  lobe 
usually  enveloping  and  capable  of  great  expansion  over  edge 
of  shell.  Upper  edge  with  finely  branched  tongue-shaped  pa- 
pillae that  alternate  with  shorter  papillae.  Lower  edge  with 
closely  spaced  tongue-shaped  papillae  of  lesser  size.  Mantle 
lobe  and  foot  side  black  except  for  lower  mantle  edge,  on 
which  branched  papillae  are  bright  yellow-orange.  The  bright 
yellow  color  of  the  lower  edge  makes  a very  conspicuous  ring 
that  encircles  the  animal,  a coloration  not  present  in  other 
species.  Determination  of  living  specimens  is  readily  made 
on  this  feature  alone;  in  preserved  specimens  it  remains  light- 
er colored  that  the  rest  of  the  mantle  lobe. 

Habitat.  Lowermost  intertidal  zone  in  partially  protected 
areas,  and  in  the  sublittoral  to  depths  of  5 m,  living  exposed 
on  the  upper  surfaces  of  rocks.  At  Antofagasta  I found  that 
the  sublittoral  population  occurred  along  with  a few  speci- 
mens of  F.  cumingi  and  still  fewer  F.  maxima.  The  rocky 
substratum  there  looked  barren  from  urchin  grazing,  but  all 
Fissurella  shells  had  a thick  algal  mat. 

Distribution.  Chiclayo,  Peru  (6°47'  S)  (Pena,  1950),  to  Rio 
Bio-bio,  Concepcion  Province,  Chile  (36°48'  S)  (LACM  75- 
35,  McLean).  I have  found  it  abundantly  as  far  north  as  Isla 
Guanape,  Peru  (8°32'  S),  and  at  all  stations  throughout  the 
range.  It  undoubtedly  occurs  further  south  than  Concepcion, 
but  I did  not  find  it  at  Mehuin,  near  Valdivia,  or  at  Guabun 
at  the  northwest  tip  of  Isla  de  Chiloe. 

Number  of  Lots  Examined.  82  (LACM  26,  AMNH  18, 
ANSP  8,  MACN  7,  MNHN  10,  USNM  13). 

Taxonomic  History.  The  normal  color  form  of  F.  lati- 
marginata has  been  understood  by  authors.  Its  synonyms  are 
based  upon  color  forms  with  the  rayed  pattern,  except  for 
Pilsbry’s  F.  punctatissima,  which  was  based  upon  a gerontic 
specimen. 

Abundance  and  Use.  Fissurella  latimarginata  is  common 
throughout  its  range.  It  occurs  widely  in  Peru,  where  few  of 
the  other  species  are  known.  It  is  one  of  the  most  important 
food  species,  more  so  than  any  other  in  northern  Chile.  The 
largest  specimens  are  taken  by  divers.  Its  common  name  is 
the  “lapa  viuda,”  which  means  widow,  for  its  black  aspect. 

Characteristics  and  Variability.  Fissurella  latimarginata 
is  characterized  by  its  generally  uniform  gray  to  reddish- 
brown  color,  fine  but  persistent  ribs,  and  tapered  anterior 
end.  The  lateral  white  rays  that  characterize  juvenile  shells 
of  so  many  of  the  species  are  likely  to  persist  through  later 
growth  stages  in  this  species.  The  broad,  dark  internal  border 


Figures  64  through  71.  Fissurella  latimarginata  Sowerby,  1835.  Mature  shells.  (64)  3-5  m,  Isla  Guanape,  Peru.  LACM  74-3,  41.9  x 27.1 
x 1 1.7  mm.  (65)  Iquique,  Chile.  LACM  64-16,  87.9  x 64.2  x 20.7  mm.  (66)  Los  Molles,  Aconcagua  Province,  Chile.  LACM  75-28,  34.8  x 
25.5  x 9.0  mm.  (67)  Los  Molles,  Aconcagua  Province,  Chile.  LACM  75-29,  116.5  x 85.5  x 37.5  mm.  (68)  Lectotype,  F.  latimarginata 
Sowerby.  Valparaiso  or  Iquique,  Chile.  BMNH  197572,  80.2  x 55.0  x 18.7  mm.  (69)  Syntype,  F.  bella  Reeve.  Cape  Horn  (probable  error). 
BMNH  197567,  56.7  x 37.4  x 12.7  mm.  (70)  Lectotype,  F.  galericulum  Reeve.  Locality  unknown.  BMNH  1976138,  33.0  x 21.5  x 9.5  mm. 
(71)  Holotype,  F.  punctatissima  Pilsbry.  Valparaiso,  Chile.  ANSP  50262,  88  x 66  x 22  mm. 


28  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  29 


Figures  72  through  79.  Fissurella  latimarginata  Sowerby,  1835.  Preserved  specimens,  mantle  edge,  juvenile  shells,  radula,  and  cut  shell.  (72) 
Living  specimen,  showing  light  colored  lower  mantle  edge  against  dark  foot  side.  Antofagasta,  Chile.  LACM  75-20.  (73)  Mantle  lobe.  Isla 
Guanape,  Peru.  LACM  74-2,  length,  24  mm.  (74)  Preserved  specimen  with  epizoic  mytilids  Semimytilus  algosus  on  shell.  1-4  m,  Isla  San 
Lorenzo,  Peru.  LACM  74-24,  66.2  x 42.5  x 22.0  mm.  (75)  Juvenile  shell.  Los  Colorados,  Antofagasta  Province,  Chile.  LACM  75-19,  16.0  x 
10.5  x 3.9  mm.  (76)  Juvenile  shell.  Iquique,  Chile.  LACM  64-16,  13.8  x 3.5  x 3.6  mm.  (77)  Radula,  air-dried.  Antofagasta,  Chile.  LACM 
75-20,  width  of  ribbon  3.4  mm,  shell  length  85.0  mm.  (78)  Head  of  preserved  specimen,  showing  mantle  lobe  and  mouth.  Isla  Guanape,  Peru. 
LACM  74-2,  shell  length  70.0  mm.  (79)  Cut  shell,  also  showing  algal  mat.  Los  Molles,  Aconcagua  Province,  Chile.  LACM  75-29,  length  of 
cut  edge  42  mm. 

is  characteristic,  but  it  is  a feature  shared  with  other  species. 

Living  specimens  are  always  recognizable  by  the  bright  yel- 
low lower  edge  to  the  mantle  lobe.  Largest  specimens  seen 
were  from  central  Chile;  smaller  specimens  occur  at  both 
extremes  of  the  range.  Shell  height  varies  extensively  within 
populations.  Specimens  with  patterns  of  rays  like  those  of  F. 
cumingi  (Fig.  61)  are  uncommon  but  occur  throughout  the 
range.  There  are  no  geographic  differences  of  sufficient  im- 
portance to  warrant  recognition  of  subspecies. 

Affinity  and  Comparisons.  Fissurella  latimarginata  most 
resembles  F.  cumingi,  having  in  common  the  size,  propor- 
tions, and  tapered  anterior  end.  The  unusual  pattern  of  rays 
in  variant  specimens  of  F.  latimarginata  is  similar  to  the 
normal  pattern  in  F.  cumingi.  The  sculpture  of  F.  latimar- 
ginata is  finer,  sharper,  and  less  beaded  than  that  of  F.  cum- 
ingi; the  rare  color  form  of  F.  latimarginata  may  always  be 
distinguished  on  this  difference  in  sculpture.  Shells  of  F.  la- 


timarginata may  resemble  those  of  F.  nigra  but  are  more 
sharply  ribbed,  have  a reddish  rather  than  gray  interior  bor- 
der, and  lack  the  inwardly  tapered,  white  bordered  foramen 
of  F.  nigra.  A similar  elaboration  of  the  tongue-shaped  pa- 
pillae of  the  mantle  lobe  is  known  only  in  F.  bridgesii.  Both 
have  broad  margins  and  are  dark  colored  with  an  incipient 
tendency  to  produce  faintly  rayed  variants.  Fissurella  lati- 
marginata may  always  be  distinguished  by  its  fine,  sharp 
ribbing,  which  is  lacking  in  F.  bridgesii. 

Synonymy  and  Types.  There  are  six  syntypes  of  F.  lati- 
marginata Sowerby,  1835,  described  from  “Valparaiso  and 
Iquique,”  Chile,  four  from  one  lot,  BMNH  197572,  lengths 
80.2,  73.8,  61.7,  and  51.1  mm,  and  two  in  BMNH  197573, 
lengths  56.3  and  27.3  mm.  Both  original  mounting  boards 
are  penned  with  “Valparaiso  and  Iquique,”  so  it  is  not  known 
from  which  of  the  two  localities  each  shell  originated.  The 
shells  are  clean  and  uniformly  dark  red,  faintly,  or  not  at  all, 


30  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


showing  the  lateral  white  rays.  The  largest  specimen  is  figured 
here  (Fig.  68)  and  designated  the  lectotype;  Dell  (1971)  fig- 
ured the  73.8  mm  specimen. 

Type  material  of  F.  biradiata  Sowerby  was  not  received 
on  loan  from  the  British  Museum.  The  type  locality  is  Val- 
paraiso, Chile.  Figure  23  of  the  “Conchological  Illustrations” 
shows  a finely  ribbed  shell  41  mm  long  (presumed  life-size) 
in  which  the  only  color  pattern  consists  of  the  two  lateral 
white  rays.  The  type  figure  can  therefore  be  relegated  to  the 
normal  form  of  F.  latimarginata  without  question.  Figure 
52  of  the  “Conchological  Illustrations”  is  stated  to  be  a “var.” 
of  F.  biradiata  from  Iquique;  this  specimen  is  the  one  figured 
by  Reeve  (1849)  and  Dell  (1971).  Reeve’s  coloration  shows 
it  to  be  reddish  brown  and  faintly  rayed.  Until  I can  examine 
the  sculpture  of  that  specimen  I am  unable  to  decide  whether 
it  is  F.  cumingi  or  the  rare,  rayed  form  of  F.  latimarginata, 
but  the  question  is  of  minor  importance  because  the  type  of 
F.  biradiata  is  clearly  recognizable  as  a specimen  of  F.  la- 
timarginata. 

Fissurella  bel/a  Reeve,  1849,  supposedly  from  Cape  Horn, 
is  represented  by  two  syntypes,  BMNH  197567,  lengths  56.7 
and  42.5  mm.  The  smaller  specimen  was  figured  by  Reeve 
(here  designated  the  lectotype)  and  the  larger  one  by  Dell 
(1971)  and  refigured  here  (Fig.  69).  No  author  has  recognized 
a species  based  on  this  name.  Dell  suggested  an  affinity  with 
F.  pulchra,  but  the  absence  of  flecking  rules  that  out.  The 
narrowed  anterior  end  and  rayed  pattern  is  shared  only  with 
the  rare  color  form  of  F.  latimarginata  and  normal  F.  cum- 
ingi. The  shells  appear  to  be  acid-cleaned,  the  fine  radial  ribs 
are  like  those  of  F.  latimarginata  rather  than  F.  cumingi. 
The  margin  in  both  shells  is  narrow,  suggesting  a degree  of 
maturity  that  can  be  matched  with  F.  latimarginata  at  ex- 
tremes of  its  distribution,  but  not  with  similarly  sized  F. 
cumingi.  I therefore  identify  the  syntypes  as  the  rare  color 
form  of  F.  latimarginata.  The  Cape  Horn  locality  is  therefore 
erroneous. 

There  are  three  syntypes  of  F.  galericulum  Reeve,  1850, 
BMNH  1976138,  lengths  33.0,  3 1 .0,  and  30.5  mm,  described 
with  unknown  locality.  The  largest  (here  designated  the  lec- 
totype) is  illustrated  here  (Fig.  70).  It  is  clearly  the  normal 
white-rayed  young  stage  of  F.  latimarginata  and  has  been  so 
recognized  by  previous  authors. 

Fissurella  punctatissima  Pilsbry,  1890,  from  Valparaiso, 
Chile,  was  considered  by  Pilsbry  to  differ  from  F.  latimar- 
ginata in  being  more  elevated  and  having  a narrower  internal 
margin.  Shell  proportions  of  the  holotype  (ANSP  50262,  Fig. 
7 1 ) and  paratype  (ANSP  61923)  are  within  the  normal  range 
of  variation  and  the  narrow  margin  is  that  of  mature  ex- 
amples of  the  species.  The  interior  pitting,  which  suggested 
the  name,  is  not  unusual  in  large  shells.  Dell’s  records  (1971) 
under  this  name  are  based  upon  specimens  of  F.  nigra. 

Fissurella  cumingi  Reeve,  1 849 

Figures  80-94 

Fissurella  cumingi  Reeve,  1849,  pi.  3,  fig.  17;  Hupe,  1854: 
238;  Sowerby  II,  1 862: 1 87,  figs.  5,  1 32;  Carcelles  and  Wil- 
liamson, 1951:256;  Dell,  1971:184,  pi.  3,  figs.  15,  16;  Ra- 
mirez-Boehme,  1974:32  [key]. 


Fissurella  latimarginata  var.  cumingi,  Pilsbry,  1890:154,  pi. 

30,  fig.  1;  Riveros-Zuniga,  1951:28. 

Fissurella  stellata  Reeve,  1850,  pi.  12,  fig.  80;  Hupe,  1854: 

245;  Sowerby  II,  1862:187,  fig.  82;  Pilsbry,  1890:148,  pi. 

32,  fig.  32;  Dali,  1909:242;  Riveros-Zuniga,  1951:1  13,  fig. 

23  [looks  like  F.  peruviana ];  Dell,  1971:193,  pi.  4,  figs.  7, 

8 [not  5,  6];  Ramirez-Boehme,  1974:30  [key]. 

Shell.  Large  (80  to  100  mm  mature  length);  height  me- 
dium; outline  oval,  tapered  anteriorly;  sides  slightly  raised. 
Sculpture  of  fine  to  medium  strength  radial  ribs,  nodulous 
or  beaded  in  early  stages  along  growth  increments;  primary 
ribs  only  slightly  more  prominent  than  secondary  ribs.  Ground 
color  dark  yellowish  gray,  patterned  with  dark  rays  of  red- 
dish-purple,  anteriormost  rays  often  split.  Margin  very  broad 
and  flat  in  young  shells,  showing  pattern  of  rays  across  the 
full  width.  Cut  shells  not  showing  zoning  of  calcitic  layer, 
pigmentation  of  rays  of  equal  intensity  throughout  layer.  Fo- 
ramen elongate  and  tripartite  in  young  shells,  elongate-oval 
in  mature  shells. 

Juvenile  Shell.  Oval,  conical,  with  straight  slopes,  earliest 
area  white,  reddish  rays  emerging  with  edges  darkly  outlined 
as  if  split;  lateral  white-rayed  area  prominent.  At  length  of 
4 mm  scattered  brown  flecks  may  be  present,  forming  zigzag 
pattern  in  some  specimens;  by  this  size  ribs  have  become 
prominent,  secondary  ribs  nearly  the  size  of  primary  ribs. 

Mantle  and  Foot.  Nearly  retractable  in  shell.  Cephalic  ten- 
tacles yellowish  on  inner  surface.  Side  of  foot  with  strongly 
developed  tubercles;  mantle  lobe  broad,  upper  edge  with 
finely  branched  papillae,  lower  edge  with  more  prominent 
papillae,  banded  in  light  and  dark  to  correspond  to  rays  on 
shell.  Foot  and  mantle  colored  purplish  red,  lighter  or  tending 
toward  pink  in  individuals  with  lightly  pigmented  shells.  The 
reddish  coloration  is  characteristic  and  differs  from  that  of 
all  other  species. 

Habitat.  Lowermost  intertidal  zone  and  immediate  sub- 
tidal  zone  to  1 5 m,  living  on  the  upper  surfaces  of  rocks.  At 
low  tide  chiefly  in  deep  tide  pools  that  occur  in  surf  exposed 
areas.  I obtained  living  specimens  from  the  municipal  market 
at  Iquique,  evidently  taken  by  divers.  Living  specimens  were 
collected  in  lesser  numbers  along  with  F.  latimarginata  at  5 
m on  a rocky  bottom  at  Antofagasta.  Large  specimens  were 
found  in  shell  piles  at  Los  Molles  (32°14'  S).  Intertidally 
occurring  specimens  were  common  at  Montemar  and  Me- 
huin. 

Distribution.  Matarani,  Peru  (17°00'  S)  (AMNH  150892, 
B.  Marco),  to  Mehuin,  Valdivia  Province,  Chile  (39°23'  S) 
(LACM  75-36,  McLean).  The  distribution  probably  extends 
farther  to  the  north  and  south.  Its  distribution  is  more  south- 
ern than  that  of  F.  latimarginata  and  F.  maxima,  both  of 
which  extend  much  farther  to  the  north  in  Peru,  but  seem 
not  to  be  represented  at  Mehuin,  where  F.  cumingi  is  com- 
mon. 

Number  of  Lots  Examined.  4 1 (LACM  16,  AMNH  6,  ANSP 
1,  MACN  3,  NMHN  10,  USNM  5). 

Taxonomic  History.  Fissurella  cumingi  has  not  hitherto 
been  understood  and  recognized  as  a separate  species.  Pre- 
vious authors  have  repeated  the  early  published  descriptions 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  31 


Figures  80  through  87.  Fissurella  cumingi  Reeve,  1849.  Mature  shells.  (80)  17  m,  Huayquique,  Tarapaca  Province,  Chile.  LACM  90799, 
32.5  x 20.5  x 1 1.2  mm.  (81)  Los  Molles,  Aconcagua  Province,  Chile.  LACM  75-28,  50.3  x 34.2  x 10.5  mm.  (82)  Same  locality  (shell  pile). 
LACM  75-29,  96.3  x 61.1  x 22.8  mm.  (83)  Lectotype,  F.  cumingi  Reeve.  Valparaiso,  Chile.  BMNH  197565,  67.2  x 49.1  x 15.2  mm.  (84) 
Montemar,  Valparaiso  Province,  Chile.  LACM  75-30,  49.9  x 34.4  x 12.8  mm.  (85)  Lectotype,  F.  stellata  Reeve,  1850.  1 1-22  m,  Valparaiso, 
Chile.  BMNH  197549,  16.2  x 10.8  x 3.8  mm.  (86)  Montemar,  Valparaiso  Province,  Chile.  LACM  75-30,  30.0  x 20.4  x 7.9  mm.  (87)  75- 
36,  Mehuin,  Valdivia  Province,  Chile.  LACM  75-36,  61.2  x 42.2  x 17.7  mm. 


and  have  considered  it  to  be  a color  form  or  “variety”  of  F. 
latimarginata.  Confusion  over  the  identity  of  F.  biradiata 
Reeve  (a  synonym  of  F.  latimarginata)  and  F.  bridgesii  Reeve 
(a  good  species)  has  also  contributed  to  the  difficulty.  Cleaned 
specimens  found  in  old  collections  have  usually  been  mis- 
identified  as  F.  maxima;  specimens  covered  with  the  algal 
mat  are  easily  mistaken  for  F.  latimarginata. 

Abundance  and  Use.  Fissurella  cumingi  is  not  as  common 
as  F.  maxima  and  F.  latimarginata.  but  large  specimens  can 


be  found  by  divers,  and  the  species  is  exploited  for  food  in 
central  and  northern  Chile.  I have  seen  it  in  the  market  at 
Iquique,  and  it  is  a major  component  in  the  shell  piles  in 
central  Chile.  The  common  name  is  “lapa  frutilla,”  meaning 
strawberry,  which  aptly  describes  the  color  of  the  mantle  and 
foot  sides. 

Characteristics  and  Variability.  The  most  characteristic 
features  of  F.  cumingi  are  its  tapered  front,  consistent  color 
pattern  of  dark  ribs  on  a relatively  dark  ground,  beaded  ribs, 


32  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


Figures  88  through  94.  Fissurella  cumingi  Reeve,  1849.  Radula,  cut  shell,  juvenile  shells,  living  and  preserved  bodies,  mantle  lobe.  (88) 
Radular  ribbon,  air-dried.  2-5  m,  Antofagasta,  Chile.  LACM  75-20,  width  of  ribbon  2.8  mm.  (89)  Cut  shell.  Los  Molles,  Aconcagua  Province, 
Chile.  LACM  75-29,  length  of  cut  edge  38.0  mm.  (90)  Juvenile  shell.  Los  Molles,  Aconcagua  Province,  Chile.  LACM  75-28,  12.8  x 7.8  x 
3.4  mm.  (91)  Juvenile  shell.  Same  locality.  LACM  75-28,  9.4  x 5.5  x 2.5  mm.  (92)  Ventral  view  of  living  specimen,  head  at  right.  Montemar, 
Valparaiso  Province,  Chile.  LACM  75-30.  (93)  Preserved  specimen.  2-5  m,  Antofagasta,  Chile.  LACM  75-20,  shell  length  73.9  mm.  (94) 
Mantle  lobe.  Same  locality.  LACM  75-20,  length  14  mm. 


and  the  unique  reddish  color  of  the  mantle  and  foot.  Vari- 
ation in  the  species  is  minimal.  The  chief  variable  feature 
seems  to  be  the  intensity  of  ground  color,  which  may  be  pale 
to  dark  yellow.  The  rayed  pattern  is  consistent;  none  have 
been  seen  that  lack  it.  Specimens  received  from  M.  Bretos 
from  the  subtidal  mussel  beds  at  Iquique  (Fig.  80)  are  small, 
elevated,  and  have  narrow  margins.  Those  from  the  south- 
ernmost locality  collected  (Mehuin,  Chile)  appeared  stunted, 
and  many  had  narrow  shells.  Flowever,  there  seem  to  be  no 
geographic  differences  of  sufficient  importance  to  recognize 
subspecies. 

Affinity  and  Comparisons.  Fissurella  cumingi  has  certain 
features  in  common  with  both  F.  maxima  and  F.  latimar- 
ginata,  suggesting  that  it  is  related  to  both.  Its  color  pattern 
combines  the  rayed  pattern  of  F.  maxima,  with  the  addition 
of  some  of  the  darker  ground  color  of  F.  latimarginata.  Its 
margin  shows  the  pattern  of  rays  across  the  full,  flat  width, 
unlike  the  solid  margin  of  F.  latimarginata  and  the  rounded, 
crenulate  and  zoned  margin  of  F.  maxima.  It  has  the  same 
size  and  proportions  as  F.  latimarginata,  but  specimens  with 
the  algal  mat  may  be  recognized  by  the  rayed  pattern  of  the 


margin.  The  normal,  rayed  form  of  F.  cumingi  may  be  dis- 
tinguished from  the  unusual  rayed  form  of  F.  latimarginata 
by  the  strength  of  the  ribs;  they  are  fine  and  sharp  in  F. 
latimarginata,  coarser  and  beaded  in  F.  cumingi.  Its  outline 
is  more  tapered  and  its  ribbing  finer  than  that  of  either  F. 
picta  lata  or  F.  costata;  it  does  not  have  the  more  distinct 
primary  ribbing  of  either  of  these  species.  Juvenile  shells  of 
F.  cumingi  have  a characteristic  color  pattern  of  split  rays; 
they  do  not  have  the  rounded  margin  of  F.  maxima  nor  the 
primary  rays  of  F.  costata  or  F.  picta  lata.  The  reddish  color 
of  the  animal  is  so  different  from  that  of  any  other  species 
that  it  may  be  recognized  by  this  feature. 

Synonymy  and  Types.  There  are  6 syntypes  of  F.  cumingi 
Reeve,  1849,  from  Quintero,  Chile,  BMNH  197565,  lengths 
77.0,  67.2,  62.5,  57.1,  32.7,  and  22.5  mm.  Reeve  (1849)  and 
Dell  (1971)  figured  the  67.2  mm  shell;  Dell’s  interior  view 
is  of  the  77  mm  specimen.  The  67.3  mm  specimen  is  here 
figured  and  designated  the  lectotype  (Fig.  83). 

Fissurella  stellata  Reeve,  1850,  was  based  on  small  spec- 
imens from  “Valparaiso,  attached  to  dead  shells  at  a depth 
of  from  six  to  twelve  fathoms.”  There  are  four  syntypes, 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  33 


BMNH  197549,  lengths  16.9,  16.2,  15.0,  and  14.5  mm.  The 
16.2  mm  specimen  is  illustrated  and  designated  the  lectotype 
(Fig.  85).  These  specimens  are  clearly  the  juveniles  of  F. 
cumingi;  they  are  red-rayed,  the  ribbing  of  medium  strength 
and  nodular,  the  primary  and  secondary  ribs  nearly  equiv- 
alent. The  original  specified  depth  is  probably  accurate;  the 
species  is  known  from  the  shallow  sublittoral.  Other  authors 
have  guessed  incorrectly  with  respect  to  this  name;  Riveros- 
Zufiiga  (1951)  figured  the  brown  and  white  rayed  form  of  F. 
peruviana  as  F.  stellata.  Dell  (1971)  mixed  the  figure  numbers 
on  his  plate  4:  his  figure  of  a syntype  of  F.  stellata  should  be 
figs.  7,  8,  rather  than  5,  6. 

Fissurella  costata  Lesson,  1831 

Figures  95-108 

Fissurella  rudis  Deshayes,  1830:134;  Deshayes  in  Lamarck, 
1836:61;  Orbigny,  1841:474  [under  F.  costata].  Not  Pa- 
tella rudis  Roeding,  1798  [=  F.  nodosa  Bom,  1778], 
Fissurella  costata  Lesson,  1831:41;  Sowerby,  1835b:4,  fig. 
28;  Orbigny,  1841:474;  Reeve,  1849,  pi.  2,  fig.  14;  Hupe, 
1854:243;  Philippi,  1860:181;  Sowerby  II,  1862:187,  figs. 
15,  205;  Pilsbry,  1890:148,  pi.  30,  fig.  10,  pi.  35,  fig.  11; 
Dali,  1909:177,  241;  Ziegenhom  and  Thiem,  1925:14,  pi. 
2,  fig.  12;  Carcelles  and  Williamson,  1951:255;  Riveros- 
Zuniga,  1951:108,  fig.  21;  Dell,  1971:183,  pi.  4,  figs.  7,  8; 
Marincovich,  1973:16,  fig.  28;  Ramirez-Boehme,  1974:31 
[key]. 

Fissurella  chilensis  Sowerby,  1835a:  124;  Sowerby,  1835b:3, 
fig.  36;  Orbigny,  1841:474  [under  F.  costata], 

Fissurella  costata  var.  rubra  Ziegenhom  and  Thiem,  1925: 
14,  pi.  2,  figs.  13a,  b. 

Shell.  Medium  sized  (50  to  80  mm  mature  length),  low  to 
medium;  outline  elongate  in  young  shells,  broadly  oval  in 
mature  shells;  base  of  shell  resting  flat  in  one  plane.  Sculpture 
of  distinctly  raised  narrow  ribs  that  crenulate  margin;  inter- 
spaces narrower  than  ribs.  Primary  ribs  more  prominent  in 
young  shells,  but  in  large  shells  not  differing  from  secondary 
ribs.  Ground  color  light  yellowish  gray;  rays  gray,  often  be- 
coming faint  in  later  growth  stages.  Margin  broad,  flat,  show- 
ing gray  rays  across  full  width  but  more  distinctly  at  outer 
edge.  Calcitic  layer  of  cut  shells  not  zoned;  rays  showing 
through  full  width.  Foramen  exceptionally  small,  elongate 
and  tripartite  in  young  shells,  elongate-oval  in  mature  shells. 

Juvenile  Shell.  Conical  at  earliest  stage,  becoming  flattened 
and  elongate;  primary  ribs  light  yellow,  strongly  elevated, 
becoming  nodular  after  shell  reaches  length  of  5 mm;  earliest 
rays  reddish,  changing  to  black  by  shell  length  of  4 mm;  black 
rays  fill  interpsaces  between  primary  ribs.  With  growth,  sec- 
ondary ribs  appear  between  primary  ribs  and  black  rays  be- 
come gray  or  disappear  altogether. 

Mantle  and  Foot.  Fully  retractable  in  shell;  shell  margin 
not  raised  and  mantle  not  projecting  in  living  specimens. 
Cephalic  tentacles  dark  on  outer  side,  yellowish  on  inside 
and  at  tips.  Mantle  lobe  very  narrow,  papillae  of  both  edges 
small  and  width  little  branching.  Side  of  foot  light  pinkish 
brown,  surface  marbled.  Projecting  tubercles  are  not  readily 
apparent  in  living  specimens  but  visible  in  preserved  spec- 


imens. Elaboration  of  the  mantle  lobes  is  the  least  pro- 
nounced in  this  species. 

Habitat.  In  northern  Chile  at  Iquique  and  Antofagasta 
Fissurella  costata  occurs  on  vertical  surfaces  of  surf-exposed 
rock  walls  at  the  low  tide  line,  near  the  holdfasts  of  the  large 
brown  alga  Lessonia,  where  its  habitat  is  not  shared  by  other 
species  of  Fissurella.  In  central  Chile  I found  it  more  abun- 
dantly. At  Los  Molles  and  Montemar  it  also  occurs  on  hor- 
izontal surfaces  in  less  exposed  areas;  here  it  shares  the  hab- 
itat with  other  species.  The  shell  margin  fits  the  contours  of 
a habitual  site  of  attachment;  when  exposed  at  low  tide  the 
shell  margin  is  in  tight  contact  with  the  substrate.  Unlike  the 
other  species,  individuals  do  not  move  when  touched  by 
Fleliaster;  the  small  foramen  may  protect  them  from  access 
by  this  seastar.  The  shell  also  seems  suited  to  resist  removal 
by  the  clingfish  Sicyases  because  the  margin  is  less  raised 
that  that  of  other  species,  which  are  more  subject  to  predation 
by  Sicyases.  Specimens  are  rarely  seen  with  attached  Scurria 
parasitica. 

Distribution.  Punta  Pichalo,  Tarapaca  Province  (19°36'  S) 
(AMNH  137232,  J.  Bird),  to  Guabun,  Isla  de  Chiloe,  Chile 
(41°50'  S)  (LACM  75-40,  McLean).  Dali’s  (1909)  record  from 
Mollendo,  Peru,  was  based  on  a specimen  of  F.  maxima 
(USNM  27743).  However,  the  distribution  of  this  species 
may  extend  farther  to  the  north  into  Peru.  In  my  collecting 
only  the  northwestern  tip  of  Isla  de  Chiloe  was  sampled  at 
Guabun;  the  southern  limit  of  the  species  is  therefore  not 
certain.  It  is  apparent,  however,  that  this  species  has  a more 
southern  center  of  distribution  than  most  species  of  the  Pe- 
ruvian Faunal  Province. 

Number  of  Lots  Examined.  70  (LACM  23,  AMNH  15, 
ANSP  3,  MACM  9,  MNHN  12,  USNM  8). 

Taxonomic  History.  Fissurella  costata  has  been  reasonably 
well  understood  by  recent  authors. 

Abundance  and  Use.  Fissurella  costata  is  moderately  com- 
mon throughout  its  range.  In  northern  Chile  it  is  little  used 
for  food  because  its  habitat  is  so  exposed  to  surf  that  it  is 
usually  inaccessible  to  shore  collectors.  Beach-worn  shells 
are  common  along  the  shore  but  fresh  shells  are  seldom  seen 
in  the  shell  piles.  The  common  name  is  “lapa  senorita,” 
because  of  its  resemblance  to  Scurria  viridu/a,  which  is  known 
as  the  “senorita.” 

Characteristics  and  Variability.  Fissurella  costata  is  char- 
acterized by  its  relatively  small  foramen,  which  is  broadly 
oval  when  mature  and  narrow  in  young  stages,  its  relatively 
low  height,  and  its  consistent  color  pattern  of  gray  rays  on  a 
yellow  ground.  Its  mantle  lobe  is  very  narrow,  and  the  foot 
color  is  light  pinkish  brown.  Variations  in  height  and  outline 
are  minor;  color  variation  results  from  rays  that  either  persist 
or  fade  in  later  stages.  There  are  no  geographic  differences 
worthy  of  note.  The  low  variability  of  this  species  is  corre- 
lated with  its  rather  restricted  habitat  on  surf-exposed  rocks 
in  the  lower  intertidal  zone. 

Affinity  and  Comparisons.  Fissurella  costata  seems  to  be 
most  closely  related  to,  and  difficult  to  distinguish  from,  F. 
picta  lata.  Both  have  similar  color  patterns,  a similar,  broadly 
oval  outline,  strong  ribs,  and  primary  ribs  that  remain  strong. 
Differences  are  that  mantle  and  foot  colors  are  lighter  in  F. 


34  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


Figures  95  through  101.  Fissurella  costata  Lesson,  1831.  Mature  shells.  (95)  Punta  Jara,  Antofagasta  Province,  Chile.  LACM  75-18,  72.3  x 
59.0  x 20.4  mm.  (96)  Los  Molles,  Aconcagua  Province,  Chile.  LACM  75-28,  66.8  x 54.5  x 23.3  mm.  (97)  Islota  Concon,  Valparaiso  Province, 
Chile.  LACM  75-31,  43.3  x 31.8  x 9.9  mm.  (98)  Lectotype,  F.  rudis  Deshayes.  Paita,  Peru  (probable  error).  MNHNP,  69.1  x 61.1  x 24.7 
mm.  (99)  Rio  Bio-bio,  Concepcion  Province,  Chile.  LACM  75-35,  57.7  x 56.4  x 15.3  mm  (beach  shell).  (100)  Mehuin,  Valdivia  Province, 
Chile.  LACM  33-2,  33.2  x 25.7  x 9.4  mm.  (101)  Ancud,  Chiloe  Province,  Chile.  LACM  62-62,  35.5  x 25.5  x 7.9  mm. 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  35 


Figures  102  through  108.  Fissurella  costata  Lesson,  1931.  Living  specimens,  juvenile  shells,  radula,  and  cut  shell.  (102)  Ventral  view,  living 
specimen.  Punta  Jara,  Antofagasta  Province,  Chile.  LACM  75-18.  (103)  Living  specimens  in  place.  Los  Molles,  Aconcagua  Province,  Chile. 
LACM  75-28.  (104)  Juvenile  specimen.  Same  locality.  LACM  75-28,  14.2  x 7.9  x 3.1  mm.  (105)  Juvenile  specimen  (dead  shell).  Bahia 
Herradura,  Coquimbo  Province,  Chile.  LACM  75-25,  10.9  x 7.3  x 2.4  mm.  (106)  Radula  of  small  specimen.  Montemar,  Valparaiso  Province, 
Chile.  LACM  75-30,  width  of  field  0.8  mm,  shell  length  24.5  mm.  (107)  Mantle  lobe.  Punta  Jara,  Antofagasta  Province,  Chile.  LACM  75- 
18,  length  10  mm.  (108)  Cut  shell.  Los  Molles,  Aconcagua  Province,  Chile.  LACM  75-28,  length  of  cut  edge  31  mm. 


costata;  F.  costata  is  somewhat  lower  than  F.  picta  lata,  al- 
though the  extremes  of  variation  seem  to  overlap;  F.  costata 
does  not  have  the  tendency  to  reddish  rays  nor  the  penciled 
pattern  of  the  rays  of  F.  picta  lata.  Juveniles  of  F.  costata 
are  more  elongate,  flatter,  and  have  more  nodulous  primary 
ribs  than  those  of  F.  picta  lata.  In  northern  Chile  where  F. 
picta  lata  does  not  occur,  there  is  no  difficulty  in  recognizing 
F.  costata  as  the  species  that  is  the  most  broadly  oval  and 
has  the  smallest  foramen. 

Synonymy  and  Types.  Four  syntypes  of  F.  rudis  Deshayes, 
1830,  have  been  examined,  received  on  loan  from  the  Paris 
Museum.  The  lot  is  accompanied  by  a cardboard  mount;  the 
shells  are  75.8,  69. 1,  53.6,  and  5 1.9  mm  in  length.  The  largest 
of  these  shells  is  the  polished  specimen  mentioned  in  the 
original  description.  The  original  dimensions  were  given  as 


55  mm  length  and  43  mm  width;  one  of  the  specimens  is 
53.6  mm  long  and  42.8  mm  wide.  The  69.1  mm  shell  is 
designated  the  lectotype  (Fig.  98).  All  four  specimens  are 
typical  and  agree  with  the  thorough  and  accurate  original 
description.  The  original  locality  was  Paita,  Peru:  “This  shell 
was  first  given  to  us  by  our  friend  Lesson,  who  found  it  at 
Paita,  Peru;  then  later  we  have  found  it  commercially,  like- 
wise from  Peru.”  [Translation.]  Later,  Deshayes  in  Lamarck 
(1836)  changed  the  locality  to  “Habite  le  Chile.”  The  actual 
occurrence  of  the  species  in  Peru  is  uncertain,  and  Paita  is 
north  of  the  known  occurrence  of  any  of  the  Peruvian  species 
of  Fissurella.  This  name  is  preoccupied  by  Patella  rudis  Roe- 
ding,  1798,  a synonym  of  the  Caribbean  F.  nodosa  (Bom, 
1778). 

Type  material  of  Fissurella  costata  Lesson,  1831,  has  never 


36  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


been  illustrated,  and  specimens  are  not  in  the  collection  of 
the  Paris  Museum,  where  some  of  Lesson’s  types  are  known 
(Bouchet,  personal  communication).  Lesson’s  description  is 
insufficient  to  distinguish  the  species  from  F.  picta  lata , which 
is  closely  related  and  also  abundant  in  the  vicinity  of  Tal- 
cahuano,  Chile,  the  type  locality.  Lesson  described  the  animal 
as  blackish  and  the  shell  margin  as  bluish  with  stains  of  red, 
features  that  would  more  appropriately  apply  to  F.  picta  lata. 
However,  the  species  is  well  known  under  the  name  F.  costata 
and  no  purpose  would  be  served  in  rejecting  this  name,  even 
though  there  is  doubt  as  to  its  identity. 

Type  material  for  F.  chilensis  Sowerby,  1 835,  has  not  been 
located.  It  came  from  “Valparaiso,  found  on  rocks  in  exposed 
situations  at  low  water.”  The  habitat  is  accurately  described; 
F.  costata  is  much  more  common  than  F.  picta  lata  at  Val- 
paraiso, and  the  original  illustration  shows  a densely  ribbed 
shell  with  a very  small  foramen.  Its  identity  is  certain,  in 
agreement  with  treatment  by  previous  authors. 

Type  material  of  Fissurella  costata  var.  rubra  Ziegenhom 
and  Thiem,  1925,  has  not  been  located.  The  specimen  came 
from  Coquimbo,  Chile,  and  measured  60  x 48  x 15  mm, 
distinguished  from  the  typical  form  in  having  reddish  rays. 
Judging  from  the  illustration,  it  could  be  either  F.  costata  or 
F.  picta  lata,  although  I have  not  seen  reddish  rayed  speci- 
mens of  F.  costata.  Lateral  profile  of  the  figure  is  about  right 
for  F.  costata;  for  F.  picta  lata  it  would  be  at  the  low  extreme 
of  variation;  the  locality  is  reasonable  for  F.  costata;  to  my 
knowledge  F.  picta  lata  does  not  occur  north  of  Valparaiso, 
where  it  is  uncommon.  I therefore  favor  retaining  this  taxon 
in  the  synonymy  of  F.  costata,  admitting  that  the  other  al- 
ternative is  a possibility. 

Fissurella  picta  ( Gmelin,  1791) 

Figures  109-146 

Fissurella  picta  is  here  considered  to  have  two  geographic 
subspecies:  F.  picta  picta  in  the  Magellanic  region  of  Chile 
and  F.  picta  lata  in  south-central  Chile. 

Synonymy  for  F.  picta  picta: 

Patella  picta  Gmelin,  1791:3729. 

Fissurella  picta,  Lamarck,  1822:10;  Deshayes,  1830:131; 
Sowerby,  1 835b:  1 , figs.  4,  26;  Lamarck,  1836:559;  Orbi- 
gny,  1841:472;  Reeve,  1849,  pi.  1,  fig.  6;  Hupe,  1854:237; 
Sowerby  II,  1862:186,  figs.  10,  11,  35;  Watson,  1886:33; 
Rochebrune  and  Mabille,  1889:70;  Pilsbry,  1890:144,  pi. 
45,  figs.  9-11;  Melvill  and  Standen,  1898:102;  Strebel, 
1907:83,  pi.  2,  fig.  22;  Melvill  and  Standen,  1907:98;  Stre- 
bel, 1908:79;  Dali,  1909:242;  Melvill  and  Standen,  1914: 
115;  Ziegenhom  and  Thiem,  1925:6,  pi.  1,  figs.  1-4;  Car- 
celles,  1950:51;  Powell,  1951:85;  Carcelles  and  William- 
son, 195 1:254;  Riveros-Zuniga,  1951:96,  fig.  15;  Dell,  1971: 
191;  Ramirez-Boehme,  1974:31  [key]. 

Fissurella  atrata  Reeve,  1850,  pi.  11,  fig.  73;  Sowerby  II, 
1862:186,  fig.  71;  Pilsbry,  1890:147,  pi.  34,  fig.  59  [under 
F.  philippiana\\  Dell,  1971:190,  pi.  3,  fig.  7 [under  F.  phi- 
lippiana ]. 

Fissurella  muricata  Reeve,  1850,  pi.  14,  fig.  103;  Sowerby 


II,  1862:106,  pi.  4,  fig.  68;  Pilsbry,  1890:156,  pi.  39,  fig. 

5. 

Synonymy  for  F.  picta  lata: 

Fissurella  lata  Sowerby,  1835a:  124;  Sowerby,  1835b:3,  fig. 

63;  Reeve,  1849,  pi.  1,  fig.  5;  Hupe,  1854:243;  Sowerby 

II,  1 862: 1 87,  fig.  1 3;  Pilsbry,  1 890: 1 47,  pi.  3 1 , figs.  18,19; 

Dali,  1909:241;  Ziegenhom  and  Thiem,  1925:13,  pi.  1 , fig. 

1 1;  Carcelles  and  Williamson,  1951:255;  Riveros-Zuniga, 

1951:107;  Dell,  1971:187,  pi.  4,  figs.  12-14;  Ramirez- 

Boehme,  1974:31  [key]. 

Fissurella  navidensis  Ramirez-Boehme,  1974:17,  31  [key]. 

Shell  (F.  picta  picta).  Medium  large  (65-95  mm  mature 
length),  moderately  elevated;  outline  elongate  oval,  sides  of 
shell  slightly  raised.  Sculpture  of  sharp,  narrow  radial  ribs; 
primary  ribs  remaining  stronger  than  secondary  and  tertiary 
ribs;  occasional  specimens  with  weak  ribs.  Ground  color  white, 
sometimes  gray,  rayed  with  black  and  white;  strongest  ribs 
centered  on  both  light  and  dark  rays;  dark  rays  usually  split 
into  number  of  fine  lines  of  black,  a result  of  lack  of  pigment 
in  grooves  between  fine  ribs.  Rays  faint  on  light  ground  in 
some,  or  black  on  gray  ground  and  with  rays  interrupted  to 
produce  concentric  patterns  of  banding.  Margin  broad  and 
flat,  showing  penciled  pattern  of  rays.  Cut  shells  show  that 
pigment  of  rays  extends  through  calcitic  layer.  Foramen  elon- 
gate in  juvenile  shells,  elongate  to  oval  in  mature  shells. 

Shell  ( F . picta  lata).  Medium  large  (50-80  mm  mature 
length),  height  moderately  to  strongly  elevated;  outline  elon- 
gate-oval to  oval,  sides  of  shell  slightly  raised.  Sculpture  of 
strong  radial  ribs;  primary  ribs  remaining  strong  at  all  growth 
stages.  Ground  color  yellowish  white,  rayed  with  yellowish 
purple;  strongest  ribs  centered  on  both  light  and  dark  rays; 
dark  rays  usually  split  into  number  of  fine  lines  coinciding 
with  secondary  and  tertiary  ribs;  pigment  tending  to  be  absent 
in  grooves  between  ribs.  Color  pattern  fairly  uniform,  with 
dark  rays  on  lighter  ground.  Margin  broad  and  flat  in  young 
shells,  sharply  defined,  gray,  showing  pattern  of  rays  and 
penciled  pattern.  Cut  shells  showing  pigment  of  rays  through- 
out calcitic  layer.  Foramen  elongate  in  young  shells,  oval  in 
mature  shells. 

Juvenile  Shell.  Juveniles  of  both  subspecies  are  oval  and 
high,  margin  broad;  strong  primary  ribs  coincide  with  light 
rays,  becoming  stronger  and  slightly  nodulous  in  specimens 
longer  than  5 mm.  Secondary  ribs  develop  in  interspaces 
between  primary  ribs;  dark  rays  develop  in  rib  interspaces 
and  show  some  concentric  interruptions  in  intensity. 

Mantle  and  Foot.  Body  nearly  retractable  within  shell;  ce- 
phalic tentacles  dark  on  outer  side,  reddish  on  inner  side, 
and  yellowish  at  tips.  Mantle  lobe  relatively  narrow,  banded 
to  correspond  to  pattern  of  rays;  papillae  of  both  edges  of 
mantle  lobe  moderately  developed.  Side  of  foot  marbled  with 
light  and  dark,  tubercles  lighter  tipped;  southernmost  spec- 
imens often  lighter  overall. 

Habitat.  Mid-tidal  to  lower  intertidal  zone  on  vertical  to 
horizontal  surfaces  and  on  the  sides  of  loose  boulders;  wedged 
in  crevices  under  more  exposed  conditions;  not  extending 
into  the  sublittoral  zone.  At  the  north  end  of  the  range  in  the 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  37 


Figures  109  through  115.  Fissurella  picta  lata  Sowerby,  1835.  Mature  shells.  (109)  Montemar,  Valparaiso  Province,  Chile.  LACM  75-30, 
37.9  x 27.8  x 13.4  mm.  (110)  Holotype,  F.  navidensis  Ramirez-Boehme.  Bahia  Navidad,  Santiago  Province,  Chile.  MNHN  200376,  66  x 
51  x 22  mm.  (Ill)  Rio  Bio-bio,  Concepcion  Province,  Chile.  LACM  75-35,  24.8  x 17.6  x 8.2  mm.  (112)  Lectotype,  F.  lata  Sowerby.  Isla 
Santa  Maria,  Bahia  Concepcion,  Chile.  BMNH  197571,  83.4  x 64.3  x 31.6  mm.  (113)  Rio  Bio-bio,  Concepcion  Province,  Chile.  LACM  75- 

35,  37.9  x 28.7  x 8.8  mm.  (114)  Mehuin,  Valdivia  Province,  Chile.  LACM  75-36,  40.4  x 29.2  x 11.7  mm.  (115)  Same  locality.  LACM  75- 

36,  40.4  x 26.5  x 12.5  mm. 


vicinity  of  Valparaiso,  F.  picta  lata  is  uncommon;  the  only 
two  living  specimens  that  I found  occurred  on  horizontal 
surfaces  adjacent  to  F.  limbata.  At  Concepcion  and  Mehuin 
and  in  the  Strait  of  Magellan  where  F.  picta  picta  occurs, 
specimens  were  common  under  all  conditions  of  exposure. 
At  these  localities,  such  characteristic  northern  species  as  F. 
latimarginata  and  F.  maxima  were  scarce  or  missing,  sug- 
gesting a correlation  between  the  dominance  of  F.  picta  lata 
and  absence  of  competition  from  the  other  species. 

Distribution.  Islote  Concon,  Valparaiso  Province,  Chile 
(32°52'  S)  (LACM  75-31,  McLean),  to  Tierra  del  Fuego  and 
Isla  de  los  Estados,  Argentina,  probably  south  to  Cape  Horn 
and  east  to  the  Falkland  Islands.  Fossil  specimens  are  known 
from  shoreline  terraces  at  Comodora  Rivadavia,  Chubut 
Province,  Argentina  (45°52'  S)  (MCZ  28329),  but  living  spec- 


imens are  unknown  on  the  mainland  Patagonian  coast  north 
of  Tierra  del  Fuego.  The  subspecies  F.  picta  picta  occurs  in 
the  Magellanic  region  of  Chile,  extending  north  to  the  vicinity 
of  Isla  de  Chiloe.  Populations  that  occur  in  the  area  of  over- 
lap, chiefly  in  the  vicinity  of  Isla  de  Chiloe,  are  consistent 
but  may  have  features  that  make  assignment  to  either  sub- 
species arbitrary  (see  further  discussion  below). 

Number  of  Lots  Examined.  F.  picta  picta:  95  (LACM  19, 
AMNH  15,  ANSP  5,  MACN  33,  MNHN  10,  USNM  13). 
F.  picta  lata:  64  (LACM  11,  AMNH  7,  ANSP  2,  MACN  5, 
MNHN  35,  USNM  4). 

Taxonomic  History.  The  typical  form  of  Fissurella  picta 
from  the  Strait  of  Magellan  has  been  well  known  and  under- 
stood by  all  authors.  Juvenile  specimens  have  been  given  the 
name  F.  atrata  Reeve,  but  it  has  otherwise  not  been  burdened 


38  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


Figures  116  through  122.  Fissurella  picta  lata  Sowerby,  1835.  Bodies  of  living  and  preserved  specimens,  juvenile  shells,  radula,  mantle  lobe, 
and  cut  shell.  (116)  Living  specimen,  ventral  view,  head  at  right.  Rio  Bio-bio,  Concepcion  Province,  Chile.  LACM  75-35.  (117)  Juvenile  shell. 
Guabun,  Chiloe  Province,  Chile.  LACM  75-40,  12.6  x 8.6  x 3.3  mm.  (118)  Radula  of  small  specimen.  Rio  Bio-bio,  Concepcion  Province, 
Chile.  LACM  75-35,  width  of  field  0.9  mm,  shell  length  27.3  mm.  (119)  Juvenile  shell.  Same  locality.  LACM  75-35,  13.5  x 8.3  x 4.3  mm. 
(120)  Preserved  specimen.  Same  locality.  LACM  75-35,  shell  length  59.5  mm.  (121)  Mantle  lobe.  Island  off  Mehuin,  Valdivia  Province,  Chile. 
LACM  75-37,  length  8 mm.  (122)  Cut  shell.  Same  locality.  LACM  75-37,  length  of  cut  edge  32  mm. 


with  excessive  numbers  of  synonyms,  as  have  the  other  two 
common  southern  species,  F.  radiosa  and  F.  oriens.  Sow- 
erby’s  Fissurella  lata  has  not  previously  been  a well  under- 
stood taxon,  perhaps  because  it  is  rare  in  the  most  populated 
region  of  central  Chile.  It  is  here  for  the  first  time  regarded 
as  a northern  subspecies  of  the  well-known  F.  picta.  The 
original  description  of  F.  lata  included  the  remark:  “This 
species  approaches,  in  form  and  colouring,  very  near  to  Fiss. 
picta,  Lam.,’’  an  accurate  observation  not  noticed  by  sub- 
sequent authors.  In  his  discussion  of  this  taxon,  Riveros- 
Zuniga  merely  quoted  previous  authors  and  figured  what  is 
more  likely  to  be  a specimen  of  F.  costata.  However,  the 
Ziegenhom  and  Thiem  (1925)  figure  is  a good  representation 
of  F.  picta  lata. 

Abundance  and  Use.  Both  subspecies  are  large  enough  and 
common  enough,  at  least  at  Concepcion  and  to  the  south,  to 
be  important  as  a food  resource.  Numerous  specimens  were 
seen  in  shell  piles  at  Mehuin.  Moreno  et  al.  (1984)  have  given 
an  account  of  the  fishery  and  ecology  of  this  species  at  Me- 
huin. I have  no  information  on  the  utilization  of  the  species 
at  more  southern  localities. 


Characteristics  and  Variability.  The  typical  F.  picta  picta 
is  large-shelled,  with  coarse  radial  ribs  and  primary  ribs  that 
remain  strong;  the  margin  is  broad;  the  dark  colored  rays  are 
split  into  numerous  fine  lines  by  grooves  that  lack  pigment 
and  separate  the  fine  ribs.  Variation  is  extensive,  chiefly  in 
strength  of  ribbing  and  color  pattern.  Variations  in  outline 
of  the  base  are  unusual;  some  oval  shells  have  been  seen 
(Figs.  127,  128).  Sculpture  varies  from  coarse  to  nearly 
smooth.  Most  specimens  are  colored  with  gray  rays  on  a 
white  ground;  variants  with  dark  ground  color  are  common. 
There  are  frequent  concentric  bands  of  different  color  inten- 
sity, probably  representing  seasonal  changes  in  temperature 
and  food  availability. 

Fissurella  picta  lata  has  a broad  outline,  as  the  name  im- 
plies. Like  F.  picta  picta  it  is  also  characterized  as  a large- 
shelled  form  with  coarse  ribbing  and  primary  ribs  that  remain 
strong;  the  color  rays  are  also  split  into  thin  lines  by  deep 
grooves  in  which  the  color  is  lacking.  The  margin  is  broad 
at  all  growth  stages,  and  the  mantle  lobe  papillae  and  foot 
tubercles  are  developed  as  in  F.  picta  picta.  As  in  the  typical 
subspecies,  there  is  extensive  variation  in  F.  picta  lata.  Al- 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  39 


Figures  123  through  129.  Fissurella  picta  picta  Gmelin,  1791.  Mature  shells.  (123)  Pargua,  Llanquihue  Province,  Chile.  LACM  75-39,  77.2 
x 49.3  x 23. 1 mm.  (124)  Pumalin,  Chiloe  Province,  Chile.  LACM  75-41 , 24.5  x 16.9  x 6.5  mm.  (125)  Same  locality.  LACM  75-41,  59.6  x 
43.5  x 24.5  mm.  (126)  Islota  Nihuel,  Chiloe  Province,  Chile.  LACM  75-44,  39.8  x 26.4  x 16.4  mm.  (127)  Quellon,  Chiloe  Province,  Chile. 
LACM  75-45,  83.5  x 68.8  x 36.8  mm.  (128)  Pumalin,  Chiloe  Province,  Chile.  LACM  75-41,  48.4  x 39.6  x 18.2  mm.  (129)  Holotype,  F. 
muricata  Reeve.  Locality  unknown.  BMNH  1976144,  25.8  x 18.4  x 13.0  mm. 


though  most  specimens  are  elevated,  the  height  is  variable, 
and  low  forms  occur  in  some  populations  along  with  more 
elevated  specimens.  Ground  color  ranges  from  light  to  dark 
gray;  the  rays  are  usually  darker  than  the  ground  color  but 
in  some  cases  only  slightly  darker. 

The  major  difference  between  the  two  subspecies  is  that 
F.  picta  lata  has  a more  oval  outline  and  usually  is  more 
elevated  than  F.  picta  picta.  However,  these  are  variable 
features  within  the  species  as  a whole,  and  specimens  occur 
at  either  extreme  of  the  distribution  having  proportions  typ- 
ical of  the  other  extreme.  Specimens  from  the  area  of  overlap. 


however,  are  more  likely  to  have  the  intermediate  propor- 
tions. Another  difference  is  that  of  the  coloration  of  the  rays: 
Fissurella  picta  picta  has  dark  purple  to  gray  rays,  whereas 
F.  picta  lata  has  rays  that  more  clearly  show  the  purple  or 
reddish  coloration.  In  both  subspecies,  however,  the  rays 
change  color  to  reddish  if  the  shells  are  faded  by  exposure 
to  the  sun.  Shells  from  shell  piles  along  the  shore  are  much 
redder  than  those  of  live-collected  or  beach-worn  specimens. 

The  extreme  differences  noted  in  some  populations  around 
Isla  de  Chiloe  seem  to  be  correlated  with  conditions  of  ex- 
posure to  the  open  coastline  on  the  west,  or  to  a more  pro- 


40  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


Figures  130  through  135.  Fissurella  picta  picta  Gmelin,  1791.  Mature  shells.  (130)  Puerto  el  Hambre,  Magallanes  Province,  Chile.  LACM 
75-49,  58.2  x 40.4  x 18.3  mm.  (131)  Holotype,  F.  alrala  Reeve.  Locality  unknown.  BMNH  197564,  23.8  x 14.0  x 5.0  mm.  (132)  Puerto 
el  Hambre,  Magallanes  Province,  Chile.  LACM  75-49,  86.3  x 58.3  x 28.3  mm.  (133)  Same  locality.  LACM  75-49,  49.2  x 34.3  x 13.7  mm. 
(134)  Laredo  Bay,  Magallanes  Province,  Chile.  USNM  1 18235,  68.6  x 61.7  x 30.0  mm.  (135)  Fox  Bay,  Falkland  Islands.  LACM  90800, 
43.3  x 27.3  x 11.5  mm. 


tected  habitat  along  the  channels  to  the  east,  where  tidal 
extremes  are  greater  than  those  of  the  open  coast  and  water 
movement  is  produced  by  tidal  currents.  At  Guabun,  on  the 
northwest,  exposed  side  of  the  island  (41°50'  S,  74°02'  W), 
the  typical  purple-rayed,  oval,  elevated  form  of  F.  picta  lata 
occurs.  Not  far  away  at  Pargua,  on  the  mainland  side  of  the 
Canal  de  Chacao  (41°47'  S,  73°28'  W),  I found  the  gray-rayed, 
low,  elongate  form  typical  of  F.  picta  picta  (Fig.  123).  At 
Pumalin  on  the  mainland  opposite  the  southern  tip  of  Isla 
de  Chiloe  (42°42'  S,  72°52'  W),  the  population  was  inter- 
mediate between  the  two  extremes,  more  oval  than  typical 
for  F.  picta  picta,  but  having  no  trace  of  the  reddish  rays 
(Fig.  125).  At  Isla  Laitec  off  the  southeast  end  of  Isla  de 
Chiloe  (43°  1 2'  S,  7 3°36'  W),  shells  were  proportionately  more 
elongate,  typical  of  F.  picta  picta.  Based  on  these  four  pop- 
ulations, I consider  that  F.  picta  lata  occurs  south  along  the 
outer  coast  of  Isla  de  Chiloe  and  F.  picta  picta  occurs  to  the 
north  along  the  inner  side  of  Chiloe  Island.  I do  not  know, 
however,  whether  populations  from  the  exposed  sides  of  is- 


lands to  the  south  of  Chiloe  Island  would  also  agree  with  F. 
picta  lata. 

Affinity  and  Comparisons.  Fissurella  picta  picta  most  re- 
sembles F.  radiosa,  a smaller-shelled  species  that  is  similar 
in  overall  proportion,  has  sculpture  of  strong  ribbing  with 
persistently  strong  primary  ribs,  and  has  a similar  color  pat- 
tern. However,  F.  picta  picta  reaches  nearly  twice  the  size  of 
F.  radiosa,  has  a much  broader  margin  at  all  growth  stages, 
and  has  a more  centrally  placed  foramen.  The  penciled  effect 
of  the  dark  rays  of  F.  picta  picta  is  not  seen  in  F.  radiosa. 
Fissurella  picta  picta  does  not  occur  sympatrically  with  any 
of  the  large  species  of  northern  Chile.  It  differs  from  all  of 
them  sufficiently  to  require  no  comparison. 

The  subspecies  F.  picta  lata  does  occur  sympatrically  with 
many  of  the  other  species.  The  latter  subspecies  is  most  closely 
related  to,  and  difficult  to  distinguish  from,  F.  costata.  Fis- 
surella picta  lata  tends  to  be  higher,  more  darkly  rayed,  and 
to  have  sculpture  with  stronger  primary  ribs  and  a slightly 
larger  foramen  than  that  of  F.  costata.  Separation  of  the  two 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  41 


Figures  136  through  146.  Fissurella  picta  picta  Gmelin,  1791.  Living  and  preserved  specimens,  radulae,  juvenile  shells,  and  cut  shells.  (136) 
Ventral  view  of  living  specimen.  Pumalin,  Chiloe  Province,  Chile.  LACM  75-41.  (137)  Mantle  lobe.  Puerto  el  Hambre,  Magallanes  Province, 
Chile.  LACM  75-49,  length  12  mm.  (138)  Air-dried  radula.  Isla  Laitec,  Chiloe  Province,  Chile.  LACM  75-47,  width  of  ribbon  1.9  mm,  shell 
length  54.3  mm.  (139)  Preserved  specimen.  Isla  de  Los  Estados,  Argentina.  LACM  71-284,  shell  length  71.8  mm.  (140)  Juvenile  specimen. 
Pumalin,  Chiloe  Province,  Chile.  LACM  75-41,  10.9  x 6.7  x 3.6  mm.  (141)  Juvenile  specimen.  Puerto  Espanol,  Bahia  Aguirre,  Tierra  del 
Fuego,  Argentina.  LACM  73-67,  17.4  x 10.6  x 3.6  mm.  (142)  SEM  photo  of  radula.  Width  of  field  1.0  mm.  (143)  Radula  of  small  specimen. 
Isla  Laitec,  Chiloe  Province,  Chile.  LACM  75-47,  width  of  field  0.7  mm,  shell  length  27.9  mm.  (144)  Cut  shell.  Pumalin,  Gulfo  Corcovado, 
Chile.  LACM  75-41,  length  of  cut  37  mm.  (145)  Cut  shell.  Puerto  el  Hambre,  Magallanes  Province,  Chile.  LACM  75-49,  length  of  cut  33.2 
mm.  (146)  Living  specimen  attached  to  substrate.  Same  locality.  LACM  75-49. 


42  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


species  on  shell  characters  may  be  difficult  and  sometimes 
entirely  arbitrary,  as  the  range  of  variation  in  the  two  species 
seems  to  overlap.  It  is  easy  to  distinguish  the  living  animals, 
however.  The  foot  and  mantle  lobe  of  F.  picta  lata  are  gray, 
those  of  F.  cost  at  a a pale  pinkish  brown. 

Synonymy  and  Types.  Fissurel/a  picta  was  known  to  pre- 
Linnaean  authors.  It  was  probably  the  first  of  the  Chilean 
species  to  reach  Europe  because  it  is  so  common  in  the  Strait 
of  Magellan.  Gmelin  (1791)  is  credited  with  the  name;  his 
knowledge  of  it  came  from  figures  in  several  previous  non- 
binomial works.  Lamarck  and  Deshayes  referred  to  it  as  the 
“Fissurelle  de  Magellan.”  Deshayes  credited  the  name  picta 
to  Lamarck,  but  Sowerby  II  (1862)  and  Pilsbry  (1890)  cor- 
rectly credited  the  authorship  to  Gmelin.  Type  material  is 
unknown. 

The  holotype  of  Fissurella  atrata  Reeve,  1850,  is  BMNH 
197564  (Fig.  131),  length  23.8  mm,  locality  unknown.  It  is 
a small,  dark-colored  specimen  of  F.  picta,  easily  recognized 
by  its  relatively  broad,  dark  inner  margin. 

The  holotype  of  F.  muricata  Reeve  is  BMNH  1976144, 
locality  unknown  (Fig.  129).  It  is  a small  dark  specimen,  the 
margin  sufficiently  broad  to  relate  it  to  F.  picta  rather  than 
F.  radiosa.  Because  the  interior  has  not  previously  been  fig- 
ured, the  dark  margin  has  been  missed,  which  explains  why 
no  author  has  related  it  to  any  of  the  Peruvian-Magellanic 
species. 

There  are  two  syntypes  of  F.  lata  Sowerby,  BMNH  197571, 
from  Isla  Santa  Maria,  Bahia  Concepcion,  Chile,  dimensions 
83.4  mm  x 64.3  x 31.6  mm,  and  77.0  x 61.3  x 25.5  mm. 
The  larger  specimen,  figured  by  Dell  (1971),  is  figured  here 
and  designated  the  lectotype  (Fig.  1 1 2).  Both  specimens  are 
reddish  rayed  and  clearly  show  the  strong  primary  ribs  in 
the  light  interspaces  between  the  rays. 

The  holotype  of  F.  navidensis  Ramirez-Boehme,  1974, 
from  Bahia  Navidad,  Santiago  Province,  Chile  (33°56'  S, 
71°52'  W),  MNHN  200376  (Fig.  1 10),  is  a worn  specimen 
of  F.  picta  lata,  with  which  it  was  not  compared.  Its  similarity 
to  F.  picta  was  noted  by  its  author,  however,  and  the  differ- 
ences described  are  those  that  are  here  used  to  distinguish 
the  two  subspecies  of  F.  picta. 

Fissurella  radiosa  Lesson,  1831 

Figures  147-175 

Fissurella  radiosa  is  here  considered  to  have  two  geographic 
subspecies:  F.  radiosa  radiosa  in  the  Magellanic  region  of 
Chile  and  Argentina,  and  F.  radiosa  tixierae  in  the  Golfo 
San  Matias  and  Peninsula  Valdez  region  of  Argentina. 

Synonymy  for  F.  radiosa  radiosa: 

Fissurella  radiosa  Lesson,  1831:411;  Orbigny,  1841:473; 

Pilsbry,  1890:157;  Strebel,  1907:85,  pi.  1,  figs.  4,  5a-d,  pi. 

9,  fig.  6;  Melvill  and  Standen,  1914:115;  Carcelles,  1950: 

51;  Carcelles  and  Williamson,  1951:254;  Riveros-Zuniga, 

1951:111;  Dell,  1971:1 92;  Ramirez-Boehme,  1974:32  [key]; 

Scarabino,  1977:178,  pi.  1,  fig.  5. 

Fissurella  picta  var.  radiosa,  Ziegenhom  and  Thiem,  1925: 

8,  11,  pi.  1,  fig.  6. 

Fissurella  nigra  Philippi,  1845:60;  Philippi,  1846,  pi.  2,  fig. 

22;  Reeve,  1849,  pi.  6,  fig.  37.  Not  F.  nigra  Lesson,  1831. 


Fissurella  darwinii  Reeve,  1849,  pi.  1.  fig.  7;  Hupe,  1854: 
247;  Rochebrune  and  Mabille,  1 889:74;  Pilsbry,  1890:144, 
pi.  30,  fig.  7,  pi.  46,  figs.  15-17;  Melvill  and  Standen,  1 898: 
102;  Strebel,  1907:93;  Carcelles,  1950:51;  Carcelles  and 
Williamson,  1951:254;  Riveros-Zuniga,  1951:98,  fig.  16; 
Dell,  1971:185,  pi.  4,  fig.  4;  Ramirez-Boehme,  1974:30 
[key]. 

Fissurella  picta  var.  darwinii,  Ziegenhom  and  Thiem,  1925: 
8,  11,  pi.  1,  fig.  5. 

Fissurella  grisea  Reeve,  1849,  pi.  6,  fig.  6;  Sowerby  II,  1862: 
184,  pi.  239,  fig.  85;  Pilsbry,  1890:152,  pi.  39,  fig.  9. 
Fissurella  exquisita  Reeve,  1850,  pi.  11,  fig.  74;  Hupe,  1854: 
246;  Sowerby  II,  1862:186,  figs.  32,  128;  Rochebrune  and 
Mabille,  1889:74;  Strebel,  1908:78,  pi.  5,  figs.  74a-c;  Car- 
celles and  Williamson,  1951:256;  Riveros-Zuniga,  1951: 
112,  fig.  22;  Metivier,  1969:115,  fig.  IB  [radula];  Dell, 
1971:185,  pi.  5,  figs.  1,  3;  Ramirez-Boehme,  1974:30. 
Fissurella  philippiana  Reeve,  1850,  errata  page;  Sowerby  II, 
1862:186,  fig.  30;  Pilsbry,  1890:146,  pi.  33,  fig.  40,  pi.  58, 
figs.  24-26;  Dali,  1909:242;  Ziegenhom  and  Thiem,  1925: 
13,  pi.  1,  figs.  10a,  10b;  Carcelles  and  Williamson,  1951: 
253;  Riveros-Zuniga,  1951:106,  fig.  19;  Dell,  1971:190. 
Fissurella  philippii  Hupe,  1854:245  (new  name  for  F.  nigra 
Philippi,  not  Lesson). 

Fissurella  polygona  Sowerby  II,  1862,  fig.  177  (not  fig.  137); 
Pilsbry,  1890:148,  pi.  60,  fig.  84;  Melvill  and  Standen, 
1898:102;  Melvill  and  Standen,  1914:115;  Carcelles  and 
Williamson,  1951:254;  Dell,  1971:192,  pi.  4,  figs.  9-1 1. 
Fissurella  dozei  Rochebrune  and  Mabille,  1885:108;  Roche- 
brune and  Mabille,  1889:72,  pi.  5,  fig.  4;  Carcelles,  1950: 
51;  Carcelles  and  Williamson,  1951:255;  Riveros-Zuniga, 
1951:101,  fig.  17;  Dell,  1971:185. 

Synonymy  for  F.  radiosa  tixierae: 

Fissurella  tixierae  Metivier,  1969: 1 16,  fig.  1 A [radula],  pi.  1, 
figs.  1-3,  9. 

Shell  (F.  radiosa  radiosa).  Small  to  medium  sized  (40  to 
55  mm  mature  length),  low  to  moderately  elevated;  outline 
elongate  oval,  somewhat  tapered  anteriorly;  sides  of  shell 
raised.  Sculpture  of  sharply  raised,  narrow  ribs  that  crenulate 
margin;  primary  ribs  remaining  stronger  and  more  promi- 
nent at  all  growth  stages.  Ground  color  varying  from  white 
to  gray  or  black  with  gray  or  reddish  gray  rays;  the  pattern 
of  rays  frequently  interrupted  by  concentric  changes  in  color 
intensity.  Primary  ribs  coinciding  with  light  rays;  in  uni- 
formly dark  shells  primary  ribs  slightly  lighter  in  color.  Mar- 
gin relatively  narrow  at  all  growth  stages,  not  zoned;  cut  shells 
showing  ground  color  or  pattern  of  rays  of  uniform  intensity 
throughout  calcitic  layer.  Foramen  just  anterior  of  center, 
elongate  and  tripartite  at  all  growth  stages. 

Shell  (F.  radiosa  tixierae).  Small  (20  to  45  mm  mature 
length),  moderately  to  strongly  elevated;  outline  elongate  ob- 
long, tapered  anteriorly;  margin  more  or  less  in  same  plane. 
Sculpture  of  narrow  ribs  that  finely  crenulate  margin.  Ground 
color  white  to  dark  gray  or  black  with  gray  or  reddish  gray 
rays,  often  interrupted  by  concentric  changes  in  intensity. 
Primary  ribs  coinciding  with  light  rays.  Margin  narrow  at  all 
growth  stages,  showing  pattern  of  rays.  Foramen  slightly  an- 
terior of  center,  elongate  and  tripartite  at  all  growth  stages. 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  43 


Figures  147  through  160.  Fissurella  radiosa  radiosa  Lesson,  1831.  Mature  shells.  (147)  Pumalin,  Chiloe  Province,  Chile.  LACM  75-41, 47.0 
x 32.0  x 13.1  mm.  (148)  Same  locality.  LACM  75-41,  41.9  x 27.4  x 11.8  mm.  (149)  Same  locality.  LACM  75-41,  43.5  x 28.4  x 13.7  mm. 
(150)  “Syntype”  [no  standing  as  type]  F.  phdippiana  Reeve.  “Chile."  BMNH  197562,  42.0  x 22.8  x 8.6.  (151)  Lectotype,  F.  darwinii  Reeve. 
Strait  of  Magellan,  Chile.  BMNH  197563,  36.9  x 22.8  x 11.0  mm.  (152)  Lectotype,  F.  grisea  Reeve.  Locality  unknown.  BMNH  1975140, 
30.3  x 19.9  x 9.9  mm.  (153)  Lectotype,  F.  exquisita  Reeve.  Locality  uncertain.  BMNH  197561,  23.4  x 14.2  x 5.2  mm.  (154)  Puerto  el 
Hambre,  Magallanes  Province,  Chile.  LACM  75-49,  40.2  x 23.0  x 10.6  mm.  (155)  Same  locality.  LACM  75-49,  38.2  x 21.4  x 8.2  mm. 
(156)  Falkland  Islands.  USNM  368377,  46.7  x 20.1  x 9.4  mm.  (157)  Lectotype,  F.  polygona  Sowerby  II.  Falkland  Islands.  BMNH  1976151, 


44  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


Figures  161  through  166.  Fissurella  radiosa  (ixierae  Metivier,  1969.  Mature  shells.  (161)  San  Antonio  Oeste,  Rio  Negro  Province,  Argentina. 
MACN  13361,  48.3  x 31.3  x 1 4.3  mm.  (162)  Puerto  Lobos,  Chubut  Province,  Argentina.  MCZ  288334,  24.6  x 14.3  x 8.3  mm.  (163) 
Holotype,  F.  tixierae  Metivier.  Golfo  Nuevo,  Chubut  Province,  Argentina.  MNHNP,  22.5  x 13.7  x 8.0  mm  (beach  shell).  (164)  Puerto 
Madryn,  Golfo  Nuevo,  Chubut  Province  Argentina.  LACM  34858,  24.9  x 14.9  x 8.5  mm  (beach  shell).  (165)  Punta  Cracker,  Golfo  Nuevo, 
Chubut  Province,  Argentina.  LACM  78-90,  27.0  x 18.8  x 8.8  mm.  (166)  Puerto  Madryn,  Golfo  Nuevo,  Chubut  Province,  Argentina.  USNM 
152895,  25.5  x 15.6  x 10.6  mm  (beach  shell). 


Juvenile  Shell.  Elongate,  margin  narrow,  ribs  fine  and  sharp, 
primary  ribs  lighter  in  color,  secondary  and  tertiary  ribs  aris- 
ing after  shell  reaches  5 mm  in  length. 

Mantle  and  Foot.  Nearly  retractable  within  shell.  Mantle 
lobe  relatively  narrow,  banded  to  correspond  to  pattern  of 
rays;  black-shelled  individuals  also  banded.  Papillae  mod- 
erately developed,  finely  branched;  side  of  foot  dark,  tuber- 
cles with  lighter  tips. 

Habitat.  Lowermost  intertidal  zone  and  offshore  to  at  least 
20  m.  In  1975  I found  it  common  at  Pumalin  in  the  Golfo 
Corcovado  on  the  undersides  of  rocks  in  an  area  where  the 
exposure  is  limited  chiefly  to  swiftly  moving  tidal  currents; 
I also  observed  it  in  the  sublittoral  at  Isla  'Falcon.  In  the 


Strait  of  Magellan  it  was  common  at  low  tide  under  rocks  at 
Puerto  Hambre.  Paul  Dayton  collected  it  by  diving  at  Isla 
de  los  Estados  in  1973.  In  1978  I found  F.  radiosa  tixierae 
to  be  common  in  the  Golfo  Nuevo  and  Golfo  San  Jose, 
Argentina,  on  undersides  of  rocks  at  low  tide  and  dredged 
offshore  to  20  m. 

Distribution.  Golfo  Corcovado  on  the  east  side  of  Isla  Chi- 
loe,  Chile  (northernmost  specimens  examined  from  Pumalin, 
Chiloe  Province,  Chile,  42°42'  S,  72°52'  W,  LACM  75-41) 
to  Tierra  del  Fuego,  probably  south  to  Cape  Horn,  east  to 
the  Falkland  Islands,  and  north  in  Argentina  to  the  Golfo 
San  Matias  (northernmost  specimens  from  San  Antonio  Oeste, 
Rio  Negro  Province,  40°45'  S,  64°58'  W,  MACN  13361,  A. 


42.4  x 28.9  x 12.3  mm.  (158)  Puerto  Deseado,  Santa  Cruz  Province,  Argentina.  LACM  34851,  27.5  x 15.7  x 5.8  mm.  (159)  Puerto  San 
Julian,  Santa  Cruz  Province,  Argentina.  AMNH  182640,  45.0  x 26.4  x 12.4  mm.  (160)  Santa  Cruz  River,  Santa  Cruz  Province,  Argentina. 
ANSP  88526,  41.4  x 25.4  x 13.5  mm. 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  45 


Figures  167  through  175.  Fissurella  radiosa  radiosa  Lesson,  1831,  and  F.  radiosa  tixierae  Metivier,  1969.  Radula,  cut  shell,  mantle  lobe, 
juvenile  shell,  living  and  preserved  specimens.  Figures  167  through  172.  F.  radiosa  radiosa.  (167)  Radula.  Puerto  el  Hambre,  Magallanes 
Province,  Chile.  LACM  75-49,  width  of  field  0.8  mm,  shell  length  20.8  mm.  (168)  Cut  shell.  Pumalin,  Chiloe  Province,  Chile.  LACM  75-41, 
length  of  cut  edge  18.5  mm.  (169)  Mantle  lobe.  Fuerte  Bulnes,  Magallanes  Province,  Chile.  LACM  75-48,  length  1 1 mm.  (170)  Juvenile  shell. 
Puerto  el  Hambre,  Magallanes  Province,  Chile.  LACM  75-49,  13.8  x 7.8  x 3.3  mm.  (171)  Juvenile  shell.  Pumalin,  Chiloe  Province,  Chile. 
LACM  75-41,  10.0  x 6.0  x 2.7  mm.  (172)  Preserved  specimen.  Fuerte  Bulnes,  Magallanes  Province,  Chile.  LACM  75-48,  shell  length  56.5 
mm.  Figures  173  through  175.  Fissurella  radiosa  tixierae.  (173)  Living  specimen  attached  to  substrate.  Punta  Cracker,  Golfo  Nuevo,  Chubut 
Province,  Argentina.  (174)  Preserved  specimen.  Punta  Ninfas,  Golfo  Nuevo,  Chubut  Province,  Argentina.  LACM  78-88,  shell  length  19.3 
mm.  (175)  Ventral-lateral  view  of  living  specimen.  Same  locality.  LACM  78-88. 


Carcelles).  The  subspecies  F.  radiosa  tixierae  is  characteristic 
only  of  the  Golfo  San  Matias  and  the  Golfo  Nuevo  and  Golfo 
San  Jose;  specimens  from  such  localities  as  Puerto  Deseado 
and  Puerto  San  Julian,  Santa  Cruz  Province,  Argentina,  are 
consistently  larger  and  lower  in  profile,  identified  as  F.  ra- 
diosa radiosa.  This  is  the  only  species  of  Fissurella  that  ranges 
throughout  the  Magellanic  Faunal  Province  in  both  Chile 
and  Argentina,  and  the  only  one  that  does  not  extend  into 
the  region  of  overlap  with  the  Peruvian  Faunal  Province  in 
south-central  Chile. 

Number  of  Lots  Examined.  F.  radiosa  radiosa:  55  (LACM 
14,  AMNH  5,  ANSP  1,  MACN  27,  MNHN  1,  USNM  7); 
F.  radiosa  tixierae:  28  (LACM  7,  AMNH  1,  MACN  19, 
USNM  1). 

Taxonomic  History.  Fissurella  radiosa  Lesson,  1831,  was 
not  originally  illustrated.  Most  of  the  accounts  dealing  with 
this  species  have  consisted  of  copies  and  translations  of  orig- 
inal descriptions  of  its  numerous  synonyms.  Accounts  with 


additional  observations  are  those  of  Strebel  (1907),  who  was 
the  first  to  recognize  the  species,  Ziegenhom  and  Thiem 
(1925),  and  Riveros-Zuniga  (1951),  who  recognized  a spec- 
imen under  the  name  of  F.  dozei  Rochebrune  and  Mabille. 
The  species  has  been  misidentified  as  Lucapinel/a  henseli 
(Martens,  1 900),  from  Puerto  Deseado,  Santa  Cruz  Province, 
Argentina,  by  Ringuelet  et  al.  (1962). 

The  Argentinian  subspecies  F.  radiosa  tixierae  was  de- 
scribed as  a distinct  species  by  Metivier  (1969),  based  on  a 
single  specimen.  He  also  identified  F.  exquisita  Reeve  from 
the  Golfo  Nuevo  (here  considered  a synonym  of  F.  radiosa), 
apparently  not  having  sufficient  specimens  to  realize  that  a 
single  species  is  represented  in  the  Golfo  Nuevo. 

Abundance  and  Use.  Fissurella  radiosa  is  common 
throughout  its  range.  It  is  a rather  small  form  occurring  to 
the  south  of  the  populated  regions  of  Chile;  I have  no  infor- 
mation as  to  whether  it  has  been  exploited  for  food. 

Characteristics  and  Variability.  The  shell  of  Fissurella  ra- 


46  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


diosa  radiosa  is  relatively  small  and  elongate,  with  a narrow 
dark  margin,  the  ribbing  fine  and  sharp,  and  the  primary  ribs 
evident  at  all  growth  stages.  Shell  height  varies  from  low  to 
moderately  high.  Color  variation  includes  rayed  forms  and 
some  that  are  uniformly  dark.  Changes  in  color  often  occur 
with  growth.  Some  shells  are  nearly  colorless  in  early  stages 
and  later  acquire  rays;  others  are  strongly  rayed  at  first  and 
then  lose  the  rays  entirely.  Normally  rayed  specimens  may 
have  growth  increments  that  are  uniformly  dark.  The  ribs 
can  be  very  evident  or  so  weak  that  one  can  barely  distinguish 
primary  ribs  from  secondary  ribs.  In  the  collections  at  hand 
there  seem  to  be  more  of  the  weakly  sculptured  examples 
from  the  vicinity  of  the  Strait  of  Magellan,  whereas  the  strongly 
sculptured  specimens  are  known  from  more  northern  local- 
ities in  Chile  and  Santa  Cruz  Province,  Argentina.  The  uni- 
formly dark  specimens  have  been  seen  only  at  Pumalin,  in 
the  Golfo  Corcovado,  where  they  occur  with  rayed  forms. 
The  largest  observed  specimen  of  the  typical  subspecies  is 
65  mm  in  length  (LACM  75-42,  Isla  Talcon,  Chile). 

The  large  series  of  specimens  of  F.  radiosa  tixierae  that  I 
collected  in  the  Golfo  Nuevo  and  Golfo  San  Jose  in  1978 
are  as  variable  in  color  as  those  of  the  typical  subspecies, 
including  many  that  are  uniformly  dark  (Figs.  161,  162). 
There  is  such  a preponderance  of  elevated  specimens  that 
the  separation  of  a geographic  subspecies  based  on  this  fea- 
ture is  justified.  However,  some  specimens  are  as  low  as  the 
typical  subspecies.  The  largest  specimen  observed  measures 
48.3  mm  in  length  (Fig.  161). 

Affinity  and  Comparisons.  Fissurella  radiosa  most  resem- 
bles F.  picta.  Fissurella  radiosa  is  smaller,  has  a narrower 
margin,  a more  anterior  foramen,  and  lacks  the  penciled 
pattern  of  the  rays.  Unlike  F.  picta  picta,  which  has  gray  rays 
only,  there  are  some  reddish-rayed  forms.  In  shell  mor- 
phology F.  radiosa  approaches  F.  oriens,  which  it  may  re- 
semble in  size,  shape,  and  range  of  color  possibilities,  but  it 
differs  in  having  a narrower  margin,  more  anterior  foramen, 
and  primary  ribs  that  are  raised,  slightly  nodular,  and  larger 
than  the  adjacent  ribs,  in  contrast  to  the  nearly  smooth  aspect 
of  F.  oriens.  The  papillae  of  the  mantle  fold  are  more  strongly 
developed  in  F.  radiosa  than  in  F.  oriens.  Fissurella  radiosa 
is  more  elongate  than  F.  peruviana  and  has  an  elongate  rather 
than  oval  foramen. 

Synonymy  and  Types.  Type  material  of  F.  radiosa  Lesson, 
1831,  from  the  Falkland  Islands,  has  never  been  illustrated 
and  may  not  be  extant.  It  is  not  housed  in  the  Paris  Museum 
where  some  of  Lesson’s  material  is  now  known  (Bouchet, 
personal  communication).  Lesson’s  description  stated:  “The 
ribs  are  separated  by  profound  narrow  grooves,  and  they  are 
arranged  with  three  small  ribs  between  each  pair  of  larger 
ones,  all  over.”  This  could  also  apply  to  F.  picta,  which  was 
then  well  known,  but  the  broad  margin  of  F.  picta  was  not 
mentioned;  hence,  by  elimination  we  are  left  with  F.  radiosa. 
Strebel  ( 1 907)  was  the  first  to  use  the  name  F.  radiosa  in  the 
sense  adopted  here. 

Type  material  of  F.  nigra  Philippi,  1 845,  not  Lesson,  1831, 
has  not  been  located.  The  locality  was  given  only  as  “Chile.” 
Philippi’s  illustration  and  description  indicate  that  the  shell 
was  predominantly  black,  the  young  stages  with  lighter  rays, 


the  ribbing  strong  and  unequal,  and  the  margin  narrow.  Spec- 
imens collected  at  Pumalin,  east  of  Isla  de  Chiloe,  are  a good 
match  (Fig.  149),  and  the  synonymy  with  F.  radiosa  is  cer- 
tain. Both  Reeve  and  Hupe  were  to  offer  replacement  names 
for  the  preoccupied  name  of  Philippi. 

Fissurella  darwinii  Reeve,  1849,  from  the  “Straits  of  Ma- 
galhaens,”  is  represented  by  four  syntypes,  BMNH  197563, 
lengths,  37.0,  36.9,  33.9,  and  21.3  mm.  The  smallest  spec- 
imen has  a margin  broader  than  any  in  the  three  larger  ones 
and  is  clearly  a young  F.  picta.  Dell  (1971)  figured  the  second 
largest  specimen  (36.9  mm  in  length);  this  is  here  figured  and 
designated  the  lectotype  (Fig.  151).  The  rays  are  reddish  pur- 
ple. Because  the  margin,  which  has  not  previously  been  il- 
lustrated, is  narrow,  I identify  the  type  lot  as  F.  radiosa. 
Pilsbry  (1890)  copied  Reeve’s  illustration  but  also  identified 
and  figured  a shell  with  a broader  margin  — that  figure  is  here 
identified  as  F.  picta. 

Fissurella  grisea  Reeve,  1849,  described  without  locality, 
is  represented  by  two  syntypes,  BMNH  1976140,  measuring 
30.3  and  21.1  mm  in  length.  The  largest  (Fig.  152)  has  been 
figured  by  Reeve  and  Sowerby  II  (1866)  and  is  here  desig- 
nated the  lectotype.  Authors  have  not  recognized  this  taxon. 
The  shell  is  gray  and  has  faint  dark  rays.  The  surface  of  the 
shell  has  evidently  been  treated  with  acid  and  only  a trace 
of  radial  sculpture  remains.  I tentatively  assign  it  to  the  syn- 
onymy of  F.  radiosa,  suggested  particularly  by  the  tripartite 
foramen. 

There  are  5 specimens  labeled  F.  exquisita  Reeve,  1850, 
BMNH  197561.  One  small  specimen  is  broken;  the  other 
four  measure  45.3,  37.8,  34.0,  and  23.4  mm  in  length.  The 
Reeve  locality  is  “Strait  of  Magalhaens,”  but  the  original 
label  reads  “Falkland  Islands.”  The  smallest  intact  specimen 
(Fig.  1 53)  is  a good  match  for  the  Reeve  figure  and  was  figured 
and  designated  the  lectotype  by  Dell  (1971).  The  largest  spec- 
imen was  also  figured  by  Dell.  All  specimens  are  relatively 
low,  have  a narrow  margin,  are  similarly  rayed  with  reddish 
brown,  and  clearly  show  strong  primary  ribs. 

Fissurella philippiana  Reeve,  1850,  was  a name  introduced 
on  the  errata  page  of  the  Fissurella  monograph:  “Sp.  37.  For 
F.  nigra  Philippi,  read  F.  philippiana  Reeve.”  Reeve  was 
renaming  a homonym,  although  he  did  not  explicitly  state 
this.  The  specimens  considered  syntypes  by  Dell  therefore 
have  no  standing  as  types.  There  are  four  specimens,  BMNH 
197562,  the  locality  “Chile”  in  ink  on  the  original  mounting 
board,  although  Reeve  gave  the  locality  as  “Southern  Chile.” 
The  specimens  measure  42.0,  38.6,  33.6,  and  32.3  mm  in 
length.  They  represent  the  dark  form  of  F.  radiosa  in  which 
the  primary  ribs  are  prominent  and  the  margin  narrow.  Dell 
(1971)  figured  the  38.6  mm  specimen;  the  largest  is  figured 
here  (Fig.  150).  The  “syntypes”  are  very  similar  to  those  I 
found  at  Pumalin  (Fig.  149).  Dali  (1909)  inexplicably  gave 
the  locality  as  “Concepcion,  Chile,”  and  this  has  been  re- 
peated by  subsequent  authors.  No  specimens  corresponding 
to  this  locality  have  been  found  at  the  USNM.  Concepcion 
is  well  to  the  north  of  the  known  distribution  of  F.  radiosa. 

Fissurella  philippii  Hupe,  1854,  was  proposed  as  a replace- 
ment name  for  F.  nigra  Philippi,  not  Lesson.  Hupe  was  un- 
aware of  the  similar  name  proposed  by  Reeve. 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  47 


j'ofeiA 


Figures  176  through  191.  Fissurella  oriens  fulvescens  Sowerby,  1835,  and  F.  oriens  oriens  Sowerby,  1835.  Mature  shells.  Figures  176  and 
177.  F.  oriens  fulvescens.  (176)  Lectotype,  F.  fulvescens  Sowerby.  Valparaiso,  Chile.  BMNH  197568,  39.7  x 23.0  x 6.8  mm.  (177)  Islota 
Concon,  Valparaiso  Province,  Chile.  LACM  75-31,  23.2  x 14.8  x 4.5  mm  (beach  shell).  Figures  178  through  191.  F.  oriens  oriens.  (178) 
Paralectotype,  F.  oriens  Sowerby.  Locality  uncertain.  BMNH  197575,62.4  x 35.7  x 14.8  mm.  (179)  Lectotype,  F.  mexicana  Sowerby.  Locality 
unknown.  BMNH  1944593,  38.6  x 22.2  x 8.6  mm.  (180)  Pargua,  Llanquihue  Province,  Chile.  LACM  75-39,  45.8  x 19.6  x 6.8  mm.  (181) 
Pumalin,  Chiloe  Province,  Chile.  LACM  75-41,  58.0  x 32.7  x 12.0  mm.  (182)  4-13  m,  Islota  Nihuel,  Chiloe  Province,  Chile.  LACM  75- 
43,  41.5  x 24.3  x 8.5  mm.  (183)  Same  locality.  LACM  75-42,  49.3  x 33.0  x 18.4  mm.  (184)  Holotype,  F.  oblonga  Ramirez-Boehme.  Punta 


48  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


There  are  two  syntypes  of  F.  polygona  Sowerby  II,  1862, 
BMNH  1976151,  from  the  Falkland  Islands.  The  largest 
measures  42.4  mm  in  length  and  was  figured  by  Dell  (1971). 
It  is  here  designated  the  lectotype  (Fig.  157).  The  smaller 
specimen  measures  22.2  mm  in  length  and  has  proportions 
similar  to  the  larger.  Primary  ribs  are  not  as  pronounced  as 
shown  in  the  original  figure  (copied  by  Pilsbry,  1890);  the 
rays  are  reddish  brown.  The  specimens  represent  the  broad, 
elevated  extreme  of  F.  radiosa,  which  may  be  the  typical 
form,  as  it  has  the  same  general  type  locality  as  that  of  the 
senior  synonym. 

Type  material  of  Fissurella  dozei  Rochebrune  and  Mabille, 
1885,  described  from  Santa  Cruz,  Patagonia,  was  not  located 
in  the  Paris  Museum  by  P.  Bouchet,  although  type  material 
of  two  other  Rochebrune  and  Mabille  taxa  is  known  in  the 
Paris  Museum.  The  drawing  given  by  the  authors  in  1 889  is 
a fair  rendition  of  F.  radiosa  with  well  marked  ribs,  the  figure 
showing  that  there  are  primary  ribs  that  are  slightly  more 
prominent  than  the  secondary'  ribs;  the  width  of  the  margin 
is  not  mentioned.  Puerto  Santa  Cruz,  Santa  Cruz  Province, 
Argentina  (50°0T  S)  is  north  of  the  eastern  entrance  to  the 
Strait  of  Magellan.  This  is  a region  in  which  /-.  radiosa  is 
now  known  to  be  the  only  living  species  of  Fissurella.  The 
synonymy  of  F.  dozei  with  F.  radiosa  is  therefore  reasonably 
certain. 

Fissurella  tixierae  Metivier,  1969,  was  based  on  one  spec- 
imen in  poor  condition,  MNHNP  uncataloged,  from  the  in- 
tertidal zone  in  the  Golfo  Nuevo,  Argentina  (42°56'  S,  64°24' 
W),  length  22.5  mm,  width  13.7  mm,  height  8 mm  (Fig.  163). 

Fissurella  oriens  Sowerby,  1835 

Figures  176-199 

Fissurella  oriens  is  here  considered  to  have  two  geographic 
subspecies:  F.  oriens  oriens  in  the  Magellanic  region  of  Chile 
and  F.  oriens  fulvescens  in  south-central  Chile. 

Synonymy  for  F.  oriens  oriens: 

Fissurella  oriens  Sowerby,  1835a:  124;  Sowerby,  1835b:3,  figs. 
25,  60;  Reeve,  1849,  pi.  2,  fig.  13;  Hupe,  1854:237;  Sow- 
erby II,  1862:186,  figs.  19,  20;  Pilsbry,  1890:152,  pi.  46, 
figs.  18,  19,  pi.  34,  fig.  58;  Melvill  and  Standen,  1907:97; 
Strebel,  1907:88,  pi.  1 , figs.  8-14,  pi.  2,  figs.  1 5-20;  Strebel, 
1908:78,  pi.  6,  figs.  97a,  b;  Dali,  1909:242;  Melvill  and 
Standen,  1914:114;  Powell,  1951:85;  Carcelles  and  Wil- 
liamson, 1951:254;  Riveros-Zuniga,  1951:123,  fig.  30;  Dell, 
1971:185,  pi.  5,  figs.  3,  4;  Ramirez-Boehme,  1974:30  [key]. 
Fissurella  mexicana  Sowerby,  1 835b:8,  fig.  61;  Reeve,  1849, 
pi.  6,  fig.  40;  Sowerby  II,  1862:186,  figs.  26-28;  Pilsbry, 


1890:153,  pi.  34,  fig.  60;  Melvill  and  Standen,  1898:102; 
Strebel,  1907:88  [under  F.  oriens]. 

Fissurella  australis  Philippi,  1845:61;  Philippi,  1845:142; 

Strebel,  1907:88  [under  F.  oriens], 

Fissurella  alba  Philippi,  1845:61;  Philippi,  1845:34,  pi.  1, 
fig.  4;  Hupe,  1854:247;  Rochebrune  and  Mabille,  1889:71; 
Pilsbry,  1890:292,  pi.  62,  figs.  3-5;  Strebel,  1907:94,  pi.  1, 
figs.  1-3,  pi.  2,  fig.  21;  Ziegenhom  and  Thiem,  1925:14, 
pi.  2,  figs.  14a,  14b,  15;  Carcelles,  1950:51,  pi.  1,  fig.  11; 
Carcelles  and  Williamson,  1951:254;  Riveros-Zuniga,  1951: 
100;  Dell,  1971:181;  Ramirez-Boehme,  1974:30  [key]. 
Fissurella  (Corrina)  alba  Christiaens,  1973:93,  pi.  4,  figs.  46, 
47. 

Fissurella  flavida  Philippi,  1857:165;  Pilsbry,  1 890:292;  Stre- 
bel, 1907:97;  Carcelles  and  Williamson,  1951:254;  Ri- 
veros-Zuniga,  1951:  121;  Dell,  1971:186;  Ramirez-Boehme, 
1974:30  [key], 

Fissurella  hedeia  Rochebrune  and  Mabille,  1885:109; 
Rochebrune  and  Mabille,  1889:72,  pi.  5,  fig.  3;  Carcelles, 
1950:51;  Carcelles  and  Williamson,  1951:255;  Riveros- 
Zuniga,  1951,  fig.  32;  Dell,  1971:186;  Ramirez-Boehme, 
1974:30  [key]. 

Fissurella  arenicola  Rochebrune  and  Mabille,  1885:109; 
Rochebrune  and  Mabille,  1889:73,  pi.  5,  fig.  1;  Carcelles, 
1950:51;  Carcelles  and  Williamson,  1951:255;  Riveros- 
Zuniga,  1951:124,  fig.  31;  Dell,  1971:  182;  Ramirez- 
Boehme,  1974:30  [key]. 

Fissurella  ( Carcellesia)  doellojuradoi  Perez- Farfante,  1952: 
32,  fig.  1;  Christiaens,  1973:92,  pi.  4,  fig.  45. 

Fissurella  cheullina  Ramirez-Boehme,  1974:17,  30  [key],  pi. 

1,  figs.  2a-c. 

Fissurella  oblonga  Ramirez-Boehme,  1974:18,  30  [key],  pi. 

2,  figs.  4a-c. 

Synonymy  for  F.  oriens  fulvescens: 

Fissurella  fulvescens  Sowerby,  1835a:  127;  Sowerby,  1835b: 
6,  fig.  49;  Reeve,  1849,  pi.  6,  fig.  42;  Hupe,  1854:245; 
Sowerby  II,  1862:184,  fig.  36;  Pilsbry,  1890:152,  pi.  33, 
fig.  49;  Dali,  1909:241;  Riveros-Zuniga,  1951:122,  fig.  29; 
Dell,  1971:186,  pi.  5,  fig.  6;  Ramirez-Boehme,  1974:30 
[key]. 

Shell  (F.  oriens  oriens).  Small  to  medium  sized  (mature 
length  40-70  mm),  low  to  moderately  elevated;  outline  elon- 
gate oval  to  very  elongate,  lateral  profile  variable,  ranging 
from  flat  or  with  either  ends  or  sides  raised.  Sculpture  of  fine 
radial  ribs,  ribs  very  broad  and  flat  at  margin,  ending  in 
extremely  fine  crenulations;  under  magnification  fine  con- 


Chulao,  Chiloe  Province,  Chile.  MNHN  200375,  55.5  x 31.0  x 12.2  mm.  (185)  Holotype,  F.  cheullina  Ramirez-Boehme.  Isla  Quellin, 
Llanquihue  Province,  Chile.  MNHN  200327,  43  x 23  x 9 mm.  (186)  Holotype,  F.  hedeia  Rochebrune  and  Mabille.  Punta  Arenas,  Magallanes 
Province,  Chile.  MNHNP,  29  x 24  x 7 mm.  (187)  Orange  Harbor,  Chile.  USNM  17328,  37.4  x 22.9  x 8.2  mm.  (188)  13-29  m.  Strait  of 
Magellan,  Chile.  LACM  90801,  47.3  x 29.7  x 10.6  mm.  (189)  18-27  rn.  Falkland  Islands.  USNM  368309,  23.2  x 14.7  x 5.0  mm.  (190) 
Punta  Arenas,  Magallanes  Province,  Chile.  LACM  75-50,  54.2  x 33.2  x 16.9  mm.  (191)  Holotype,  F.  arenicola  Rochebrune  and  Mabille. 
Locality  uncertain.  MNHNP,  42.2  x 26.5  x 14.2  mm. 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  49 


Figures  192  through  199.  Fissure/la  oriens  oriens  Sowerby,  1835.  Living  specimens,  radula,  mantle  lobe,  juvenile  shells,  and  cut  shell.  (192) 
Detached  living  specimens.  4-13  m,  Islota  Nihuel,  Chiloe  Province,  Chile.  LACM  75-43.  (193)  SEM  view  of  radular  ribbon.  Pargua,  Llanqihue 
Province,  Chile.  LACM  75-39,  width  of  field  0.7  mm.  (194)  Radula.  Pumalin,  Chiloe  Province,  Chile.  LACM  75-41,  width  of  field  0.5  mm. 
(195)  Mantle  lobe.  Same  locality.  LACM  75-41,  length  9 mm.  (196)  Juvenile  shell.  11-13  m,  Isla  Carlos  III,  Magallanes  Province,  Chile. 
LACM  73-70,  7.8  x 4.5  x 1.9  mm.  (197)  Juvenile  shell.  4-13  m,  Islota  Nihuel,  Chiloe  Province,  Chile.  LACM  75-43,  13.4  x 8.2  x 3.4  mm. 
(198)  Air-dned  radula.  Pumalin,  Chiloe  Province,  Chile.  LACM  75-41,  width  of  ribbon  3.0  mm,  length  of  shell  48.8  mm.  (199)  Cut  shell.  4- 
13  m,  Islota  Nihuel,  Chiloe  Province,  Chile.  LACM  75-42,  length  of  cut  edge  17  mm. 


centric  lamellae  visible  in  early  growth  stages.  Ground  color 
variable,  from  colorless  to  yellow  or  dark  red;  rays  mostly 
reddish  or  gray,  solid  or  split  into  two  or  more  broad  bands 
and  frequently  with  concentric  interruptions;  gray  rays  often 
changing  to  reddish  or  black  to  gray.  Some  specimens  com- 
pletely colorless.  Margin  relatively  narrow,  somewhat  broad- 
er in  rapidly  growing  specimens,  not  zoned,  showing  full 
pigmentation  of  rays  throughout  calcitic  layer.  Foramen 
elongate  and  tripartite  in  young  shells,  elongate-oval  in  ma- 
ture shells. 

Shells  of  the  northern  subspecies,  F.  oriens  fulvescens, 
smaller  (maximum  length  45  mm),  relatively  low,  ends  usu- 
ally elevated  relative  to  sides.  Color  consistent;  ground  color 


yellow  orange,  rays  dark  red,  rays  frequently  split  into  two 
adjacent  bands,  young  shells  speckled  with  red.  This  color- 
ation and  the  speckled  pattern  of  young  shells  is  not  matched 
by  that  in  any  specimens  of  the  typical  subspecies. 

Juvenile  Shell.  Elongate  oval,  more  conical  than  at  later 
stages,  usually  reddish  overall,  with  white  apical  area  and 
color  pattern  emerging  unevenly;  rays  arising  after  shell 
reaches  about  4 mm  in  length. 

Mantle  and  Foot.  Animal  usually  large,  not  retractable  in 
flattened  shell.  Cephalic  tentacles  reddish  brown,  yellowish 
at  tips.  Mantle  lobe  rather  thin  in  preserved  specimens,  edges 
with  finely  branched  papillae,  banded  to  correspond  to  rayed 
pattern;  side  of  foot  mottled  brown  to  pinkish  brown,  light 


50  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


in  specimens  with  colorless  shells.  Epipodial  tentacles  es- 
pecially prominent. 

Habitat.  Chiefly  sublittoral  throughout  the  range,  but  ex- 
tending up  to  the  lower  intertidal  zone  in  places  exposed  to 
currents  but  not  heavy  surf.  The  northern  subspecies  F.  ori- 
ens fulvescens  is  probably  limited  to  the  sublittoral  zone,  for 
I was  unable  to  find  living  specimens  at  low  tide.  I observed 
the  typical  subspecies  at  Mehuin  in  deep  tide  pools.  I found 
it  at  Pumalin  in  the  Golfo  Corcovado,  occurring  at  low  tide 
in  rocky  areas  free  of  sand  where  tidal  currents  were  strong; 
at  Islote  Nihuel  in  the  Golfo  Corcovado  I found  it  at  a depth 
of  10  m on  rocks  near  the  sand-rock  interface.  At  Puerto 
Hambre  in  the  Strait  of  Magellan  I found  it  at  low  water  in 
sheltered  rocky  areas.  Specimens  have  been  dredged  to  depths 
of  30  m by  the  R/V  HERO  near  Isla  de  los  Estados,  Argen- 
tina. 

Distribution.  Islote  Concon,  Valparaiso  Province,  Chile 
(32°52'S)  (LACM  75-31,  McLean),  south  toTierradel  Fuego 
and  probably  Cape  Horn,  east  to  Isla  de  los  Estados,  Argen- 
tina, and  the  Falkland  Islands.  Not  known  living  from  main- 
land Argentina.  Pleistocene  specimens  are  known  from  Co- 
modoro  Rivadavia,  Chubut  Province,  Argentina  (MCZ 
288329),  as  is  also  true  for  F.  picta.  Scarabino’s  (1977)  record 
of  the  species  from  the  Golfo  San  Matias,  Argentina,  is  prob- 
ably based  upon  specimens  of  F.  radiosa  tixierae.  Popula- 
tions from  the  northern  end  of  the  range  in  the  vicinity  of 
Valparaiso  and  south  at  least  to  Concepcion  are  of  the  sub- 
species F.  oriens fulvescens.  Those  at  Mehuin  and  to  the  south 
are  the  typical  subspecies  F.  oriens  oriens. 

Number  of  Lots  Examined.  F.  oriens  oriens:  97(LACM  34, 
AMNH  10,  ANSP  6,  MACN  28,  MNHN  6,  USNM  13);  F. 
oriens  fulvescens:  4 (LACM). 

Taxonomic  History.  Fissurella  oriens  Sowerby  has  been 
reasonably  well  understood  by  most  authors,  although  the 
extent  of  its  variability  and  the  extreme  number  of  synonyms 
has  not  generally  been  recognized.  Most  of  the  synonyms 
have  not  come  into  general  use,  with  one  exception,  that  of 
F.  alba  Philippi,  which  was  based  on  white-shelled  forms. 
The  northern  subspecies,  F.  oriens  fulvescens  Sowerby,  is 
here  recognized  for  the  first  time. 

Abundance  and  Use.  Fissurella  oriens  is  primarily  a species 
of  the  Magellanic  Faunal  Province,  where  it  is  common  at 
low  water  and  much  more  abundant  in  the  sublittoral.  Al- 
though many  shells  are  small,  it  reaches  sizes  large  enough 
to  be  used  for  food.  I have  no  information  on  the  extent  of 
its  use. 

Characteristics  and  Variability.  The  most  characteristic 
features  of  F.  oriens  are  the  relatively  small  size  and  lack  of 
strong  ribbing,  the  radial  sculpture  being  better  described  as 
consisting  of  striae  or  grooves.  It  is  one  of  the  most  variable 
species  in  proportions,  size  of  the  foramen,  and  color  pattern. 
Local  populations  tend  to  be  consistent,  with  many  similarly 
appearing  individuals.  The  ground  color  of  the  typical  sub- 
species ranges  from  colorless  to  dark  red,  but  most  commonly 
has  strong  rays  that  may  change  with  growth  from  gray  to 
reddish  and  increase  or  diminish  in  intensity.  White  shelled 
specimens  have  been  noticed  in  scattered  populations 
throughout  the  range  of  the  species.  Some  populations  have 


shells  that  are  relatively  flat,  while  in  others  the  shells  are 
more  elevated.  This  is  one  of  the  few  species  in  which  the 
variation  includes  specimens  with  either  elevated  sides  or 
elevated  ends.  The  margin  is  usually  narrow,  though  not  as 
narrow  as  that  of  F.  radiosa.  However,  some  specimens  that 
are  growing  rapidly  may  have  a relatively  broad  margin.  In 
some  gerontic  specimens,  the  foramen  may  become  very 
large  and  broadly  oval;  in  others  it  remains  narrow  and  elon- 
gate. 

Affinity  and  Comparisons.  Fissurella  oriens  most  resem- 
bles F.  radiosa,  a species  of  similar  size  in  which  there  is 
similar  variation  in  proportions  and  color.  Fissurella  oriens 
differs  chiefly  in  lacking  the  strong  ribbing  of  F.  radiosa  and 
in  having  a more  centrally  placed  foramen.  However,  some 
conspicuously  ribbed  specimens  of  F.  oriens  may  so  resemble 
weakly  ribbed  specimens  of  F.  radiosa  that  the  only  reliable 
character  to  separate  them  is  the  presence  of  primary  ribs 
larger  than  the  adjacent  ribs  on  F.  radiosa,  and  the  absence 
of  such  primary  ribs  on  F.  oriens.  Strongly  rayed  specimens 
of  F.  oriens  may  have  a superficial  resemblance  to  uneroded 
specimens  of  F.  limbata,  but  F.  oriens  lacks  the  broad  two- 
zoned  margin  that  is  the  hallmark  of  F.  limbata. 

Synonymy  and  Types.  Eleven  names  seem  to  be  referable 
to  F.  oriens,  the  large  number  of  synonyms  correlated  with 
the  high  variability  of  the  species  and  the  tendency  for  local 
populations  to  have  uniform  features. 

There  are  five  syntypes  of  F.  oriens  Sowerby,  1835,  BMNH 
197575,  lengths  69.3,  62.4,  59.6,  53.7,  and  16.7  mm.  The 
largest  specimen,  that  figured  by  Reeve  ( 1 849)  and  Dell  (1971) 
is  designated  the  lectotype;  the  second  largest  specimen,  a 
paralectotype,  is  figured  here  (Fig.  1 78).  Although  Reeve  gave 
the  locality  as  Valparaiso,  Sowerby’s  original  locality  is  “In- 
sulam  Chiloe,”  with  a “variety”  mentioned  from  Valparaiso. 
The  specimens  are  relatively  large,  rayed  in  reddish  brown, 
and  represent  the  normal,  elongate,  moderately  elevated  form, 
similar  to  what  I have  observed  from  the  vicinity  of  Isla  de 
Chiloe.  The  type  locality  should  therefore  be  limited  to  Isla 
de  Chiloe.  Sowerby’s  original  figure  in  the  “Conchological 
Illustrations”  was  of  a smaller  specimen,  probably  the  59.6 
mm  specimen. 

Sowerby  did  not  clearly  document  the  above-mentioned 
“variety”  of  F.  oriens  from  Valparaiso.  However,  in  the  same 
publication  (Sowerby,  1835a)  he  described  F.  fulvescens  from 
Valparaiso,  which  may  have  been  intended  as  the  “variety.” 
In  1975  I found  beach-worn  specimens  matching  the  type 
lot  on  cobble  beaches  in  that  vicinity  of  Chile  (Fig.  177).  The 
type  lot  consists  of  four  specimens,  BMNH  197568,  lengths 
39.7,  37.0,  33.0,  and  27.9  mm.  The  largest  specimen  (Fig. 
176)  was  figured  originally  by  Sowerby  ( 1835b),  later  by  Reeve 
(1849),  and  more  recently  by  Dell  (1971),  who  designated  it 
the  lectotype.  All  the  specimens  are  low  and  elongate,  yellow- 
orange  in  ground  color,  the  rays  reddish,  and  the  margin 
narrow.  The  specimens  are  in  good  condition  and  were  ev- 
idently collected  alive;  according  to  Sowerby,  they  were  taken 
under  rocks  on  the  shore. 

Fissurella  mexicana  Sowerby,  1835,  was  said  to  come  from 
“Real  Llejos,  Mexico,”  obviously  in  error.  There  are  four 
specimens  in  the  type  lot,  BMNH  1966493,  lengths  40.6, 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  51 


40.4,  38.6,  and  29.6,  the  smallest  specimen  broken  and  re- 
paired. They  are  elongate,  moderately  elevated,  and  rather 
thin-shelled,  well  within  the  range  of  variation  of  F.  oriens. 
The  38.6  mm  specimen  is  designated  the  lectotype  (Fig.  1 79). 

Fissurella  australis  Philippi,  1845,  from  “Fretum  Magel- 
lanicum,”  was  never  illustrated  and  I have  not  been  able  to 
locate  type  material.  Philippi  described  its  color  and  sculp- 
ture as  similar  to  that  of  a young  F.  oriens , but  thinner-shelled 
and  with  extremities  raised,  the  shell  resting  on  the  sides. 
Inasmuch  as  specimens  of  F.  oriens  with  raised  ends  and 
thin  shells  are  well  known,  the  synonymy  is  reasonably  cer- 
tain. 

Fissurella  alba  Philippi,  also  from  “Fretum  Magellani- 
cum,”  was  illustrated  subsequently  by  Philippi,  but  again, 
type  material  has  not  been  located.  A white,  finely  sculptured 
shell  was  figured,  and  Philippi  himself  indicated  that  it  could 
be  a variant  of  F.  oriens.  This  note  was,  surprisingly,  over- 
looked by  Pilsbry  (1890)  and  the  taxon  has  been  accepted 
without  question  by  subsequent  authors  and  even  made  the 
type  species  of  a subgenus  based  upon  the  character  of  weak 
ribbing.  Nevertheless,  it  is  clearly  a white-shelled  variant  of 
F.  oriens.  I have  seen  white  shells  from  numerous  stations 
at  which  rayed  forms  also  occur  (see  Fig.  188,  from  a lot  of 
six  specimens,  four  of  which  are  white  and  two  are  rayed). 

Fissurella  flavida  Philippi,  1857,  from  the  “Magellen- 
strasse,”  has  not  been  figured  and  I have  not  located  type 
material.  The  description  indicates  a shell  that  is  solid,  el- 
liptical, moderately  convex,  with  obscure  sculpture,  and  a 
moderately  broad  margin.  The  coarse  sculpture  that  char- 
acterizes F.  picta  and  F.  radiosa  were  not  mentioned.  Inas- 
much as  only  three  species  occur  in  the  region,  this  name 
can  be  relegated  to  the  synonymy  of  F.  oriens  by  eliminating 
the  other  possibilities. 

The  holotype  of  F.  hedeia  Rochebrune  and  Mabille,  1 885, 
from  Punta  Arenas  in  the  Strait  of  Magellan,  was  received 
on  loan  from  the  Paris  Museum  (Fig.  186).  Although  not 
compared  by  its  authors  to  other  species,  it  is  an  elongate, 
finely  sculptured  specimen  of  F.  oriens. 

Fissurella  arenicola  Rochebrune  and  Mabille,  1885,  also 
was  not  compared  to  other  species.  Two  specimens  in  the 
Paris  Museum  labeled  “type  et  paratype,”  were  studied.  The 
locality  inked  on  the  mounting  board  is  “Baie  Orange”;  the 
published  locality  is  “Punta- Arenas  Patagoniae,  Baie  Or- 
ange.” The  holotype  (Fig.  191)  is  a white-shelled  F.  oriens, 
42.2  mm  in  length,  very  worn  except  at  the  margin,  where 
no  traces  of  primary  ribs  are  evident.  The  paratype  is  38.2 
mm  in  length,  also  white-shelled,  but  primary  ribs  are  ap- 
parent throughout.  I therefore  identify  the  paratype  as  F. 
radiosa.  Based  on  the  holotype,  the  name  F.  arenicola  is 
placed  in  the  synonymy  of  F.  oriens. 

Fissurella  ( Carcellesia ) doellojuradoi  Perez-Farfante,  1952, 
was  based  on  a single  specimen  from  “Tierra  del  Fuego.” 
The  specimen,  35  mm  in  length,  was  borrowed  by  its  author 
from  the  Museo  Argentino  de  Ciencias  Naturales  in  Buenos 
Aires.  A new  subgenus  was  based  on  the  single  feature  of  the 
raised  ends  of  the  specimen,  but  as  discussed  above,  such 
variants  are  frequently  seen  in  F.  oriens,  with  which  it  was 


not  originally  compared.  The  synonymy  of  this  taxon  there- 
fore seems  certain. 

Fissurella  cheullina  Ramirez-Boehme,  1974,  was  based  on 
eight  specimens  from  Isla  Queullin,  Llanquihue  Province, 
Chile  (41°53'  S,  72°55'  W),  holotype  MNHN  200377  (Fig. 
185).  Fissurella  oblonga  Ramirez-Boehme,  1974,  was  based 
on  a single  specimen  from  Punta  Chulao,  Chiloe  Province, 
Chile  (42°17'  S,  72°50'  W),  holotype  MNHN  200375  (Fig. 
1 84).  Neither  taxon  was  compared  by  its  author  to  any  other 
species.  Both  were  described  as  moderately  elevated,  elon- 
gate, weakly  sculptured,  and  with  narrow  margins.  These 
specimens  are  well  within  the  range  of  variation  known  for 
F.  oriens. 

Fissurella  nigra  Lesson,  1831 

Figures  200-2 1 1 

Fissurella  nigra  Lesson,  1831:412;  Orbigny,  1841:473;  Phi- 
lippi, 1846:65,  fig.  2;  Reeve,  1849,  pi.  2,  fig.  11;  Hupe, 
1854:241;  Sowerby  II,  1862:184,  fig.  14;  Pilsbry,  1890: 
149,  pi.  35,  figs.  1,2;  Dali,  1909: 1 77,  242;  Ziegenhom  and 
Thiem,  1925:15,  pi.  2,  figs.  16a,  b;  Carcelles  and  William- 
son, 1951:255;  Riveros-Zuniga,  1951:116,  fig.  25;  Dell, 
1971:188,  pi.  3,  figs.  17-19;  Ramirez-Boehme,  1974:31 
[key], 

Fissurella  violacea  Rathke,  1833  [ex  Eschscholtz  manu- 
script]^!, pi.  23,  fig.  6;  Orbigny,  1841:473  [under  F.  ni- 
gra]-, Philippi,  1846:66,  pi.  2,  fig.  3. 

Fissurella  grandis  Sowerby,  1835a:123;  1835b:3,  fig.  48;  Or- 
bigny, 1841:473  [under  F.  nigra]-,  Philippi,  1846:65,  pi.  2, 
fig-  1. 

Shell.  Relatively  large  (70  to  1 10  mm  mature  length),  mod- 
erately elevated;  outline  elongate  oval,  base  resting  flat  or 
with  ends  slightly  raised,  rarely  with  raised  sides.  Overall 
appearance  smooth,  sculptured  with  fine  radial  ribs  that  per- 
sist to  margin.  Color  black  or  gray,  sometimes  pale  and  show- 
ing concentric  variation  in  shading;  rays  lacking  or  faint, 
consisting  of  narrow  lines  slightly  darker  than  adjacent  ground. 
Margin  broad  and  flat  in  growing  shells,  narrow  in  mature 
shells;  two-zoned,  outer  zone  black,  inner  zone  translucent 
gray;  cut  shells  showing  that  two  zones  are  nearly  equal  in 
thickness.  Foramen  anterior  to  center,  elongate  and  tripartite 
in  young  shells,  elongate-oval  in  mature  shells,  worn  or  bev- 
eled at  apex  to  reveal  aragonitic  layer,  so  that  it  always  ap- 
pears white-bordered.  Old  shells  that  continue  to  grow  by 
increasing  height  without  expanding  at  base  may  contract 
basal  area,  forming  thick  edge  and  losing  distinct  zoning  of 
margin. 

Juvenile  Shell.  Radial  ribs  fine  and  sharp;  primary  and 
secondary  ribs  appearing  early.  Whitish  in  earliest  stage, 
changing  abruptly  or  gradually  to  black,  some  showing  two 
lateral  white  rays  that  quickly  fade. 

Mantle  and  Foot.  Nearly  containable  in  shell.  Entire  ani- 
mal gray;  yellow  coloration  lacking  in  cephalic  tentacles. 
Mantle  lobe  greatly  expandable,  faintly  banded,  narrow  when 
preserved;  papillae  of  upper  edge  moderately  developed,  fine- 
ly branched;  those  of  lower  edge  nearly  equal  in  size.  Side  of 


52  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


Figures  200  through  204.  Fissurella  nigra  Lesson,  1831.  Mature  shells.  (200)  Island  off  Mehuin,  Valdivia  Province,  Chile.  LACM  75-37, 
74.7  x 50.2  x 19.5  mm.  (201)  Lectotype,  F.  grandis  Sowerby,  1835.  Locality  uncertain.  BMNH  1976143,  101.0  x 67.9  x 30.9  mm.  (202) 
Quellon,  Chiloe  Province,  Chile.  LACM  75-45,  79.5  x 51.1  x 30.1  mm.  (203)  Pumalin,  Chiloe  Province,  Chile.  LACM  75-41, 43.8  x 27.5 
x 12.4  mm.  (204)  Isla  Bertrand,  off  Isla  Navarino,  Magallanes  Province,  Chile.  NMNZ  18409,  78.9  x 56.5  x 25.4  mm. 


foot  darker  than  mantle  lobe;  tubercles  small  and  widely 
scattered.  This  is  the  only  species  with  a completely  gray 
animal. 

Habitat.  Intertidal  zone  only,  on  the  undersides  of  rocks 
in  the  mid-tidal  to  lower  intertidal  zone.  Several  large  spec- 
imens may  occur  close  together  on  the  undersides  of  large 
rocks.  Shells  are  mostly  clean,  except  for  scattered  incrus- 
tations of  barnacles,  bryozoa,  or  spirorbid  worms.  None  have 


been  found  with  attached  Scurria  parasitica,  as  would  be 
expected  if  the  habitat  were  more  exposed.  This  is  the  only 
species  occurring  under  rocks  in  the  mid-littoral  of  south 
central  and  southern  Chile;  only  juveniles  of  other  species  of 
Fissurella  occur  in  this  habitat  in  northern  Chile. 

Distribution.  Valparaiso,  Santiago  Province,  Chile  (33°02' 
S)  (USNM  48218,  Bridges),  to  Puerto  Grandi,  Isla  Bertrand, 
off  south  side  Isla  Navarino,  Chile  (5  5°  1 2'  S,  67°02'  W)  (Dell, 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  53 


Figures  205  through  211.  Fissurella  nigra  Lesson,  1831.  Living  specimens,  radulae,  juvenile  shell,  cut  shell,  and  mantle  lobe.  (205)  Living 
specimen  on  underside  of  overturned  boulder.  Island  ofTMehuin,  Valdivia  Province,  Chile.  LACM  75-37.  (206)  Radula.  Same  locality.  LACM 
75-37,  width  of  field  0.8  mm,  shell  length  26.4  mm.  (207)  Ventral  view  of  living  specimen.  Rio  Bio-bio,  Concepcion  Province,  Chile.  LACM 
75-35.  (208)  Juvenile  shell.  Island  off  Mehuin,  Valdivia  Province,  Chile.  LACM  75-37,  5.5  x 3.5  x 1.5  mm.  (209)  Air-dried  radula.  Same 
locality.  LACM  75-37,  width  of  ribbon  2.6  mm.  (210)  Cut  shell.  Same  locality.  LACM  75-37,  length  of  cut  edge  39  mm.  (211)  Mantle  lobe. 
Same  locality.  LACM  75-37,  length  10  mm. 


1971).  The  northernmost  specimen  personally  collected  is  a 
single  beach-worn  juvenile  from  Punta  el  Lacho,  Santiago 
Province,  Chile  (33°30'  S).  The  species  is  rare  in  the  vicinity 
of  Valparaiso,  if  it  now  occurs  there  at  all.  Dali's  ( 1 909)  record 
from  Callao,  Peru,  is  not  represented  by  USNM  specimens 
and  should  be  discounted.  I found  this  species  commonly  at 
Concepcion  and  Mehuin,  and  at  all  stations  in  the  vicinity 
of  Isla  de  Chiloe,  both  on  the  exposed  outer  coast  and  on 
the  eastern  side  where  water  motion  is  primarily  that  of  tidal 
currents.  It  is  evidently  uncommon  and  sporadic  in  the  south- 
ernmost region,  for  I found  no  trace  of  it  at  Punta  Arenas  or 
Puerto  Hambre  in  the  Strait  of  Magellan.  It  is  unknown  from 
the  Falkland  Islands.  In  addition  to  the  above  record  of  Dell 
(1971),  the  following  southern  records  are  known  to  me: 
USNM  1 70205,  Port  Otway,  Chile  (46°49'  S);  MACN  12491, 
Canal  Smyth  (at  western  end  of  Strait  of  Magellan);  AMNH 
173403,  Beagle  Canal  (south  side  Tierra  del  Fuego). 


Number  of  Lots  Examined.  67  (LACM  1 1,  AMNH  8,  ANSP 
3,  MACN  8,  MNHN  33,  USNM  4). 

Taxonomic  History.  Fissurella  nigra  is  an  easily  recognized 
species  that  has  been  understood  by  most  authors. 

Abundance  and  Use.  Common,  at  least  in  the  northern- 
most portion  of  the  range.  Its  intertidal  habitat  is  accessible, 
and  it  is  exploited  for  food. 

Characteristics  and  Variability.  The  most  characteristic 
features  of  F.  nigra  are  the  gray  to  black  surface,  the  rays,  if 
present,  being  faint  and  split  into  lines,  and  the  prominent 
zoning  of  the  calcitic  layer  at  the  margin,  the  outer  zone  dark 
and  the  inner  zone  a light  translucent  gray.  Variation  is  chiefly 
in  color  pattern.  Some  specimens  change  from  black  to  gray 
or  light  brown;  such  specimens  being  more  likely  to  show 
the  fine  brown  radial  pattern.  There  may  be  changes  in  color 
intensity  with  growth,  but  never  as  pronounced  as  that  which 
occurs  in  F.  picta,  F.  radiosa,  or  F.  oriens.  The  size  record 


54  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


for  the  species,  perhaps  for  the  genus  as  well,  is  length  135 
mm,  width  100  mm,  height  62  mm  (Isla  de  Chiloe,  collected 
in  1892,  MNHN). 

Affinity  and  Comparisons.  Fissurella  nigra  does  not  closely 
resemble  any  other  species.  Although  its  overall  appearance 
is  smooth,  juvenile  shells  are  sharply  ribbed,  and  there  are 
definite  primary  and  secondary  ribs.  The  presence  of  these 
ribs  in  the  juvenile  dissociates  it  from  the  smooth-shelled 
species  in  the  “group  of  F.  limbata”  in  which  early  sculpture 
consists  only  of  broad  primary  ribs.  Its  closest  affinity  is 
probably  with  F.  radiosa,  with  which  it  shares  similar  pro- 
portions, an  anteriorly  positioned  foramen,  and  the  three 
series  of  ribs,  although  the  total  number  of  ribs  is  greater  in 
F.  nigra.  The  dark  outer  zone  of  F.  nigra  is  unique.  There 
is  only  a superficial  resemblance  of  F.  nigra  to  dark  gray 
specimens  of  F.  latimarginata.  Compared  to  the  latter,  F. 
nigra  has  fine  radial  ribs,  a dark  rather  than  lighter  outer 
zone  to  the  margin,  and  a more  anterior  and  more  inwardly 
beveled  foramen. 

Synonymy  and  Types.  Type  material  of  F.  nigra  Lesson, 
1831,  is  unknown.  It  is  not  represented  in  the  Paris  Museum 
where  some  of  the  Lesson  collection  is  now  housed.  Although 
there  were  no  original  illustrations,  the  species  is  easily  rec- 
ognized from  the  description.  It  is  common  in  the  vicinity 
of  its  type  locality  “Saint-Vincent,”  now  San  Vicente  (36°43' 
S),  near  Concepcion,  Chile. 

Fissurella  violacea  Rathke,  1833,  was  described  before  the 
Lesson  work  was  known.  The  original  figure  is  clear;  the  type 
locality  is  Concepcion.  Type  material  may  be  in  Leningrad, 
where  some  of  the  Eschscholtz  and  Rathke  types  are  extant. 

Fissurella  grandis  Sowerby,  1835,  is  another  early  name 
evidently  introduced  before  Lesson’s  work  was  known.  There 
are  two  specimens  in  the  type  lot,  BMNH  1976143,  lengths, 
110.1  and  77.2  mm.  The  larger  specimen  was  figured  by 
Sowerby  (1835b),  Reeve  (1849),  and  Dell  (1971).  It  is  here 
designated  the  lectotype  (Fig.  201).  The  original  locality  was 
given  as  “Valparaiso  and  Chiloe,”  so  it  is  not  certain  whether 
both  specimens  are  from  the  same  place.  The  correct  locality 
may  be  Isla  de  Chiloe,  because  the  species  is  rare  in  the 
vicinity  of  Valparaiso. 

Group  of  Fissurella  limbata 

Relatively  large-shelled  species  in  which  the  thickness  of  the 
calcitic  layer  greatly  exceeds  that  of  the  interior  aragonitic 
layer.  Radial  sculpture  in  the  early  stage  is  either  absent  or 
consists  of  broad,  low  primary  ribs;  there  are  no  secondary 
ribs.  Mature  shells  may  retain  the  broad  primary  ribs  as  low 
undulations  or  be  entirely  smooth. 

Although  shells  of  this  group  have  the  thick  calcitic  layer 
of  the  F.  maxima  group,  they  differ  from  the  latter  in  not 
having  secondary  ribs.  Whether  this  difference  in  sculpture 
has  taxonomic  significance  should  be  tested  at  the  biochem- 
ical level.  It  is  premature  to  justify  a separation  at  the  subge- 
neric level. 

There  are  four  species  in  this  group:  F.  limbata.  F.  crassa. 
F.  bridgesii,  and  F.  pulchra.  The  latter  three  have  elongate 
shells  with  low  profiles,  the  animals  too  large  to  be  retracted 
within  the  shell.  These  species  tend  to  be  more  stenotopic 


than  those  of  the  F.  maxima  group.  Variation  in  each  species 
is  less  extreme,  which  is  correlated  with  the  relatively  few 
synonyms  in  the  group. 

The  four  species  in  the  group  are  limited  to  the  Peruvian 
Faunal  Province,  unlike  the  F.  maxima  group,  in  which  there 
are  both  Peruvian  and  Magellanic  members. 

Fissurella  limbata  Sowerby,  1835 

Figures  2 1 2-224 

Fissurella  limbata  Sowerby,  1835a:  123;  Sowerby,  1835b:3, 
figs.  42,  66,  74;  Orbigny,  1841:474;  Reeve,  1849,  pi.  2, 
figs.  10,  12;  Hupe,  1854:239;  Sowerby  II,  1862:184,  figs. 
23,  24;  Pilsbry,  1890:149,  pi.  32,  figs.  26-39;  Dali,  1909: 
242;  Ziegenhom  and  Thiem,  1925;  15,  pi.  2,  figs.  17-19, 
20a,  20b;  Riveros-Zuniga,  1951:114,  fig.  24;  Pena,  1970: 
156;  Dell,  1971:188,  pi.  5,  fig.  5;  Manncovich,  1973:18, 
fig.  30;  Ramirez-Boehme,  1974:32  [key]. 

Fissurella  limbata  var.  multilineata  Ziegenhom  and  Thiem, 
1925:16,  pi.  2,  fig.  21. 

Shell.  Medium  large  (60-90  mm  mature  length),  moder- 
ately elevated;  outline  elongate  oval;  base  usually  resting  flat, 
with  ends  or  sides  slightly  raised.  Sculpture  of  wavy,  irregular 
radial  ribs;  overall  aspect  mostly  smooth.  Color  of  uneroded 
shells  (lacking  epibiotic  Scurria ) consisting  of  yellowish  ground 
and  purple  rays,  rays  often  split.  Calcitic  layer  zoned,  con- 
sisting of  inner  layer  of  solid  dark  purple  and  translucent 
outer  layer.  Eroded  shells  (those  with  Scurria)  worn  to  white 
aragonitic  layer  near  foramen;  area  farther  away  from  fora- 
men consisting  of  deep  purple  portion  of  calcitic  layer,  with 
pattern  of  rays  evident  only  near  margin.  Margin  broad  in 
growing  shells,  dark  purple  next  to  aragonitic  interior,  outer 
edge  lighter  and  revealing  pattern  of  rays.  Cut  shells  showing 
outer  zone  about  one-fourth  thickness  of  solid  purple  zone. 
Foramen  elongate  at  all  stages,  tripartite  in  young  shells, 
constricted  in  middle  in  mature  shells. 

Juvenile  Shell.  Elongate-oval,  elevated;  primary  ribs  broad, 
weak.  Color  generally  white,  marked  with  concentric  zigzag 
purple  lines  coalescing  into  purple  rays;  inner  purple  zone  of 
calcitic  layer  apparently  lacking  in  juvenile  shells. 

Mantle  and  Foot.  Shell  edge  enveloped  by  mantle  on  at- 
tached specimens  exposed  at  low  tide,  body  retractable  with- 
in shell.  Mantle  lobe  light  gray,  appearing  narrow  in  pre- 
served specimens,  marked  with  concentric  lines  of  black,  and 
only  faintly  banded  to  correspond  to  rays  of  shell.  Papillae 
at  edges  very  small,  finely  branched.  Side  of  foot  gray;  tu- 
bercles small,  scattered.  Preserved  specimens  retaining  broad 
dark  ring  with  lighter  edge  where  the  smooth  innermost  edge 
of  mantle  lobe  is  in  contact  with  shell.  This  is  the  only  species 
in  which  a dark  zone  on  the  inner  lobe  shows  in  preserved 
specimens. 

Habitat.  Mid-intertidal  to  low-intertidal  zones  in  surf-ex- 
posed areas,  primarily  on  horizontal  and  sloping  surfaces, 
rather  than  vertical  surfaces.  This  is  the  predominant  species 
on  flat  areas  of  exposed  reefs.  I saw  no  specimens  in  the 
sublittoral  zone  at  localities  where  I dived.  Most  shells  have 
a single  Scurria  parasitica,  which  produces  a deeply  etched 
attachment  scar. 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  55 


Figures  212  through  216.  Fissurella  limbala  Sowerby,  1835.  Mature  shells.  (212)  Isla  Guanape,  Peru.  LACM  74-2,  37.0  x 22.9  x 12.1  mm. 
(213)  Shore  opposite  Isla  Santa  Maria,  Antofagasta  Province,  Chile.  LACM  75-17,  65.8  x 41.0  x 20.0  mm.  (214)  Los  Molles,  Aconcagua 
Province,  Chile.  LACM  75-28,  82.4  x 57.7  x 25.0  mm.  (215)  Paralectotype,  F.  limbata  Sowerby.  Valparaiso,  Chile.  BMNH  197581, 65.0  x 
48.5  x 1 17.8  mm.  (216)  Lectotype,  F.  limbata  Sowerby.  Valparaiso,  Chile.  BMNH  197582,  60.5  x 40.0  x 17.4  mm. 


Distribution.  Isla  Guanape,  Peru  (8°30'  S)  (LACM  74-2, 
McLean),  to  Isla  de  Chiloe,  Chile  (AMNH  155914,  O.  Ruiz). 
The  exact  locality  for  the  southern  record  is  not  known,  but 
it  was  probably  the  accessible  northwestern  tip  at  approxi- 
mately 41°50'  S.  However,  I was  unable  to  find  specimens 
in  that  vicinity  at  Guabun,  Isla  de  Chiloe.  The  species  does 
not  occur  in  the  sheltered  waters  of  the  Golfo  Corcovado  on 
the  eastern  side  of  Isla  de  Chiloe.  I discount  the  record  of 
Riveros-Zuniga  (1951)  from  Fuerte  Bulnes  in  the  Magellan 
Strait. 

Number  of  Lots  Examined.  100  (LACM  35,  AMNH  20, 
ANSP  8,  MACN  8,  MNHN  10,  USNM  19). 

Taxonomic  History.  Fissurella  limbata  has  been  correctly 
interpreted  by  most  authors. 

Abundance  and  Use.  This  species  is  common  throughout 
its  range  and  is  exploited  for  food.  Large  specimens  are  sel- 


dom seen,  because  of  the  accessibility  of  the  habitat  at  low 
tide.  It  is  called  the  “lapa  gaviota”  at  Iquique.  Gaviota  is  the 
name  for  seagull,  a predator  on  this  species. 

Characteristics  and  Variability.  The  most  characteristic 
and  unusual  feature  of  F.  limbata  is  the  zonation  of  the 
calcitic  layer,  in  which  the  pattern  of  rays  is  confined  to  the 
thin,  lighter  colored  surface  layer.  The  greater  thickness  of 
the  calcitic  layer  consists  of  the  dark  purple  inner  zone,  which 
is  much  darker  than  the  rays.  There  is  little  variation;  dif- 
ferences in  appearance  are  a result  of  patterns  of  wear  in 
which  the  outermost  rayed  layer  is  lost,  leaving  a uniform 
purple  layer.  Further  wear  results  in  complete  loss  of  the 
calcitic  layer  near  the  foramen,  which  exposes  the  white  ara- 
gonitic  layer.  Erosion  of  the  shells  is  greatly  accelerated  when 
Scurria  parasitica  is  present. 

Affinity  and  Comparisons.  Fissurella  limbata  seems  not 


56  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


WiW  220 


222 


223 


219 


217 


224 


Figures  217  through  224.  Fissurella  limbata  Sowerby,  1835.  Living  specimens,  juvenile  shells,  radula,  cut  shell,  and  mantle  lobe.  (217) 
Ventral  view  of  living  specimen.  Cumbres  Borascosas,  Tarapaca  Province,  Chile.  LACM  75-14.  (218)  Juvenile  shell.  Iquique,  Chile.  LACM 
64-14,  1 1.8  x 6.8  x 3.0  mm.  (219)  Air-dried  radula.  Montemar,  Valparaiso  Province,  Chile.  LACM  75-30,  width  of  ribbon  2.4  mm.  (220) 
Radula  of  small  specimen.  Ancon,  Lima  Province,  Peru.  LACM  74-21,  width  of  held  0.5  mm,  shell  length  18.8  mm.  (221)  Juvenile  shell. 
Iquique,  Chile.  LACM  64- 16,5.4  x 3.3  x 1.6  mm.  (222)  Living  specimens  with  mantle  extended  next  to  chiton  Enoplochiton  niger.  Iquique, 
Chile.  LACM  75-12.  (223)  Cut  shell.  Mehuin,  Valdivia  Province,  Chile.  LACM  75-36,  length  24  mm.  (224)  Mantle  lobe,  Islota  Concon, 
Valparaiso  Province,  Chile.  LACM  75-31,  length  12  mm. 


closely  related  to  any  other  species.  It  has  some  features  in 
common  with  F.  crassa— sculpture  essentially  lacking  and  a 
very  elongate  foramen.  Both  are  limited  to  the  intertidal  zone, 
although  F.  limbata  is  found  at  lower  levels  than  is  F.  crassa. 
The  margin  is  broad  and  flat,  unlike  the  upturned  margin  of 
F.  crassa.  Also,  F.  limbata  has  relatively  weak  development 
of  mantle  papillae  and  foot  tubercles,  whereas  both  of  these 
features  are  strongly  developed  in  F.  crassa.  It  probably  has 
more  in  common  with  F.  nigra,  with  which  it  shares  a rel- 
atively smooth  shell,  similar  weak  development  of  mantle 


papillae,  and  foot  tubercles.  However,  the  zoning  of  the  cal- 
citic  layer  is  reversed;  the  darkest  layer  is  at  the  surface  in 
F.  nigra.  Fissurella  oriens  has  a rayed  pattern  similar  to  that 
of  F.  limbata,  but  has  some  radial  sculpture  and  an  unzoned 
calcitic  layer. 

Synonymy  and  Types.  Two  lots  of  F.  limbata  from  the 
Cuming  Collection  in  the  British  Museum  have  been  ex- 
amined. For  each  lot  the  locality  “Valparaiso”  is  written  in 
ink  on  the  boards.  Single  specimens  from  each  lot  were  il- 
lustrated by  Reeve,  1849,  although  neither  of  the  two  spec- 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  57 


Figures  225  through  231.  Fissurella  crassa  Lamarck,  1822.  Mature  shells.  (225)  Bahia  Independencia,  lea  Province,  Peru.  AHF  375-35, 
15.6  x 9.3  x 3.5  mm.  (226)  Iquique,  Chile.  LACM  64- 16,  56.9  x 32.3  x 14.0  mm.  (227)  Holotype,  F.  clypeiformis  Sowerby . Locality  unknown. 
BMNH  40.6. 1 .9,  7 1 .4  x 43.4  x 1 5.6  mm.  (228)  Cumbres  Borascosas,  Tarapaca  Province,  Chile.  LACM  75-14,  19.8  x 11.4  x 4.6  mm.  (229) 
Los  Molles,  Aconcagua  Province,  Chile.  LACM  75-28,  92.0  x 61.0  x 27.4  mm.  (230)  Vina  del  Mar,  Valparaiso  Province,  Chile.  LACM  66- 
46,  45.5  x 19.8  x 7.0  mm.  (231)  Mehuin,  Valdivia  Province,  Chile.  LACM  75-36,  80.3  x 49.0  x 22.1  mm. 


imens  illustrated  by  Sowerby  in  the  “Conchological  Illustra- 
tions” are  included  in  these  lots.  However,  there  is  no 
mistaking  the  type  figure  of  Sowerby  (1835b,  figs.  66,  74, 
internal  and  external  view)  as  this  species.  Of  the  lots  figured 
by  Reeve,  BMNH  197582  has  six  specimens,  length,  60.5 
(anterior  end  chipped),  59.2,  49.2,  48.4,  33.6,  and  28.2.  The 
largest  was  figured  by  Reeve  (1849)  and  Dell  (1971)  and  is 
here  refigured  and  designated  the  lectotype  (Fig.  216).  All 
specimens  in  this  lot  lack  Scurria- made  scars  and  show  the 
complete  normal  color  pattern  of  the  outer  zone  of  the  calcitic 
layer.  BMNH  197581  has  four  specimens,  length  67.3,  65.0, 
63.1,  and  61.1  mm.  The  largest  was  figured  by  Reeve,  al- 
though the  scar  of  a Scurria  was  omitted;  most  of  the  rayed 
outer  zone  of  the  calcitic  layer  was  removed  by  the  Scurria, 
leaving  the  shell  dark  purple  except  for  later  growth  stages. 


The  65.0  mm  shell  is  figured  here  (Fig.  215);  it  has  a Scurria- 
made  scar  but  is  less  eroded. 

Fissurella  limbata  var.  multilineata  Ziegenhom  and  Thiem, 
1925,  was  based  on  two  shells  from  Coquimbo,  Chile,  the 
largest  16.5  mm  in  length.  The  figure  shows  the  normal  zigzag 
pattern  of  purple  lines  found  in  all  juveniles  (Figs.  218,  221); 
the  name,  therefore,  has  no  systematic  value. 

Fissurella  crassa  Lamarck,  1822 

Figures  225-237 

Fissurella  crassa  Lamarck,  1822,  6(2):  11;  Deshayes,  1830: 
134;  Sowerby,  1 835b:  1 , figs.  9,  11;  Deshayes  in  Lamarck, 
1836,  7:592;  Gray,  1839:148,  pi.  39,  fig.  9;  Orbigny,  1841: 
472;  Reeve,  1849,  pi.  1,  fig.  4;  Hupe,  1854:240;  Philippi, 


58  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


Figures  232  through  237.  Fissurella  crassa  Lamarck,  1822.  Juvenile  shell,  radula,  preserved  and  living  specimens,  cut  shell,  and  mantle  lobe. 
(232)  Juvenile  shell.  Miraflores,  Lima  Province,  Peru.  LACM  71-187,  6.9  x 3.5  x 11.6  mm.  (233)  Radula  of  small  specimen.  Cartagena, 
Santiago  Province,  Chile.  LACM  75-34,  width  of  field  1.5  mm,  shell  length  22.1  mm.  (234)  Living  animal  submerged  in  tidepool.  Iquique, 
Chile.  LACM  75-12.  (235)  Preserved  specimen.  Mehuin,  Valdivia  Province,  Chile.  LACM  75-36,  shell  length  46.9  mm.  (236)  Cut  shell.  Bahia 
Moreno,  Antofagasta  Province,  Chile.  LACM  75-16,  length  35  mm.  (237)  Mantle  edge,  Iquique,  Chile.  LACM  75-12,  length  1 1 mm. 


1860:181;  Sowerby  II,  1862:184,  figs.  16,  17;  Watson,  1886: 
32;  Pilsbry,  1890:154,  pi.  34,  figs.  51-53;  Dali,  1909:177, 
241,  pi.  24,  figs.  5,  6 [fig.  looks  like  F.  limbata\\  Ziegenhom 
and  Thiem,  1925:18,  pi.  2,  fig.  24;  Carcelles  and  William- 
son, 1951:255;  Mermod,  1950:702;  Riveros-Zuniga,  1951: 
93,  fig.  14;  Pena,  1970:156;  Dell,  1971:184;  Marineovich, 
1973:17,  fig.  27;  Ramirez-Boehme,  1974:30  [key], 
Fissurella  depressa  Lamarck,  1822,  6(2):  15;  Sowerby,  1835b: 
1 (under  F.  crassa)-,  Mermod,  1950:713  [type  lost]. 
Fissurella  clypeiformis  Sowerby,  1825,  app.,  p.  vi;  Sowerby, 
1835b:  1 [under  F.  crassa]. 

Shell.  Medium  large  (60-90  mm  mature  length),  height 
low  to  moderately  elevated,  elongate  oval  to  very  elongate, 
some  with  elevated  sides,  others  with  elevated  ends  or  with 
both  (so  that  shells  rests  on  four  comers).  Sculpture  smooth 
except  for  early  primary  ribs  that  become  broad  and  low, 
forming  wide  marginal  crenulations.  Color  caramel  brown, 
occasionally  with  faint  rays  of  darker  brown,  surface  eroded 
if  epibiotic  Scurria  is  present,  clean  and  uneroded  if  not. 
Margin  upwardly  rounded  at  all  growth  stages.  Shell  margin 
showing  light  inner  zone  and  darker  outer  zone.  Cut  shells 
also  showing  very  thin  lighter-colored  layer,  layer  not  evident 


at  edge.  Foramen  elongate  and  tripartite  in  young  shells,  very 
elongate  and  constricted  in  middle  in  mature  shells,  posterior 
portion  much  wider  and  longer  than  anterior;  foramen  in 
mature  shells  beveled  inward  at  ends.  Aragonitic  layer  of 
interior  between  callus  and  muscle  scar  pinkish  gray  through- 
out and  radially  ridged. 

Juvenile  Shell.  Sculpture  of  strong  light-colored  primary 
ribs  that  become  wide  and  low,  interspaces  wide  and  dark- 
colored,  two  lateral  white  rays  prominent.  Shell  becomes 
dark  overall  on  reaching  length  of  7 mm.  Upwardly  rounded 
margin  begins  in  earliest  stages. 

Mantle  and  Foot.  Shell  too  low  to  accommodate  large  an- 
imal within  it.  Cephalic  tentacles  brown;  mantle  lobe  broad, 
banded  in  light  and  dark,  bands  not  matched  by  shell  rays 
but  no  doubt  corresponding  to  rayed  pattern  of  early  juve- 
niles. Papillae  of  upper  edge  very  large,  those  of  lower  edge 
much  less  developed.  Side  of  foot  with  large,  bulbous,white- 
tipped  tubercles.  Papillae  project  at  both  ends  of  foramen. 

Habitat.  Mid-intertidal  zone,  in  crevices  on  rocky  reefs, 
in  surf-exposed  or  partially  protected  areas,  occurring  at  higher 
levels  than  the  other  species,  tightly  wedged  in  narrow  crev- 
ices during  low  tide.  The  upturned  margin  may  be  an  ad- 
aptation to  the  rather  cramped  position  of  the  animal  when 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  59 


exposed  at  low  tide.  Most  specimens  have  a single  Scurna 
parasitica  on  the  shell.  Bretos  (1978,  1980)  has  studied  growth 
in  this  species. 

Distribution.  Huarmey,  Peru  (10°06'  S)  (AMNH  134571), 
to  Punta  Pulga,  Isla  de  Chiloe,  Chile  (42°06'  S)  (Dell,  1971). 
Dali  ( 1 909)  cited  the  Galapagos  Islands,  Ecuador,  and  USNM 
59260  is  so  labeled,  but  this  record  is  discounted.  The  south- 
ern limit  is  uncertain.  The  species  is  known  from  the  north- 
western tip  of  Isla  de  Chiioe  but  may  extend  farther  south. 
I did  not  find  it  on  the  eastern  side  Isla  de  Chiloe,  where 
there  is  little  exposure  to  surf.  The  record  of  Riveros-Zuniga 
(1951)  from  Fuerte  Bulnes  in  the  Magellan  Strait  is  rejected. 

Number  of  Lots  Examined.  88  (LACM  31,  AMNH  23, 
ANSP  7,  MACN  7,  MNHN  10,  USNM  10). 

Taxonomic  History.  Fissurella  crassa  has  been  correctly 
interpreted  by  authors.  It  differs  sufficiently  from  all  other 
species  that  it  can  not  be  confused  with  any  of  them. 

Abundance  and  Use.  Common  throughout  its  range  and 
widely  exploited  for  food.  The  upper  intertidal  habitat  is  so 
accessible  that  large  specimens  are  seldom  seen.  It  is  known 
as  the  “lapa  de  sol,”  because  it  occurs  relatively  high,  where 
it  is  exposed  to  the  sun. 

Characteristics  and  Variability.  The  most  characteristic 
features  of  F.  crassa  are  the  uniformly  brown  color  of  the 
shell,  the  lack  of  radial  ribs  other  than  the  broad  undulations, 
the  lavender  staining  of  the  interior,  the  great  enlargement 
of  the  posterior  end  of  the  foramen,  and  of  most  importance, 
the  upturned  margin.  The  foot  also  has  the  strongest  devel- 
opment of  tubercles  in  any  of  the  species.  Variation  is  not 
extensive  and  is  limited  to  rather  minor  differences  in  height 
and  amount  of  elevation  of  the  sides  or  ends. 

Affinity  and  Comparisons.  The  closest  affinity  of  F.  crassa 
seems  to  be  with  F.  limbata.  In  both  species  the  foramen 
remains  elongate,  the  sculpture  is  undulating,  and  strong  ribs 
are  lacking.  As  in  F.  limbata  there  is  a thin  outermost  zone 
to  the  calcitic  layer  that  is  lighter  in  color.  The  upturned 
margin  of  F.  crassa  is  unique.  Although  the  margin  of  F. 
maxima  is  also  unusual,  in  that  species  only  the  junction 
between  the  margin  and  the  internal  aragonitic  layer  is  round- 
ed. 

Synonymy  and  Types.  The  original  description  of  F.  crassa 
Lamarck  mentioned  no  locality.  Mermod  (1950:702),  in  his 
report  on  types  of  Lamarck  in  the  Geneva  collection,  gave 
notes  on  a specimen  73  mm  in  length,  which  he  considered 
to  be  Lamarck’s  original. 

The  locality  for  the  very  briefly  described  F.  depressa  La- 
marck. 1822,  was  given  as  the  “Indian  Ocean.”  Sowerby 
( 1 835b:  1 ) stated  under  F.  crassa:  “Lamarck’s  F.  depressa  is 
only  a worn  fragment  of  this  species,  as  Mr.  Gray  informs 
me.”  Presumably,  Gray  had  examined  the  Lamarck  collec- 
tion. Mermod  (1950:713)  reported  that  the  type  specimen 
now  is  lost. 

The  holotype  of  F.  clypeiformis  Sowerby,  1825,  is  an  un- 
worn specimen  of  F.  crassa,  BMNH  40.6.1.9,  70.4  mm  in 
length  (Fig.  227).  It  was  described  without  locality  and  was 


placed  in  synonymy  shortly  after  publication  by  Sowerby 
( 1 835b)  himself. 

Fissurella  bridgesii  Reeve,  1 849 

Figures  238-253 

Fissurella  bridgesii  Reeve,  1849,  pi.  3,  fig.  15;  Hupe,  1854: 
238;  Philippi,  1860:180;  Sowerby  II,  1862:184,  figs.  21, 
22,  34;  Pilsbry,  1890:151,  pi.  30,  fig.  3;  Dali,  1909:241; 
Riveros-Zuniga,  1951:121,  fig.  28;  Dell,  1971:183,  pi.  3, 
figs.  8,  9;  Ramirez-Boehme,  1974:31  [key]. 

Shell.  Medium  large  (65-90  mm  mature  length),  low;  out- 
line elongate-oblong,  lateral  profile  mostly  straight,  but  some 
specimens  with  slightly  raised  sides  or  ends.  Sculpture  of  fine 
low  ribs  in  young  shells;  mature  shell  nearly  smooth,  with 
traces  of  very  broad,  irregular  radial  ribs  and  irregular  radial 
striae.  Color  grayish  or  reddish  brown,  with  faint  lighter  and 
darker  rays,  two  lateral  rays  often  more  prominent;  ground 
color  changing  gradually  to  light  gray  in  large  shells.  Margin 
broad  in  growing  shells,  flat  and  often  beveled  out  and  up 
(except  at  front  end)  so  that  its  edge  is  not  in  contact  on  a 
flat  surface;  margin  narrower  and  rounded  in  mature  shells. 
Calcitic  layer  zoned,  inner  zone  dark  reddish  brown,  changing 
to  translucent  gray  at  outer  surface,  outer  zone  usually  lighter. 
Foramen  unusually  large  and  oval  at  all  growth  stages,  broad- 
ly tripartite  in  some  and  showing  two  projections  at  sides, 
others  lacking  these  projections. 

Juvenile  Shell.  None  seen  under  20  mm  in  length;  primary 
and  secondary  ribs  weak,  nearly  equal  in  size  at  shell  length 
of  20  mm;  foramen  large  and  oval  at  this  shell  length.  Young 
shells  have  two  lateral  white  rays. 

Mantle  and  Foot.  Body  too  large  to  retract  beneath  low 
shell;  the  mantle  lobe  broad  and  thin,  enveloping  shell  edge. 
Papillae  of  upper  edge  tongue-shaped,  finely  branched,  those 
of  lower  lobe  similarly  shaped  but  smaller  and  more  nu- 
merous. Mantle  lobe  vertically  banded  with  light  and  dark 
to  correspond  to  weak  pattern  of  rays  on  shell.  Side  of  foot 
marbled;  in  preserved  specimens,  dark  tubercles  are  sur- 
rounded by  lighter  areas. 

Habitat.  Fissurella  bridgesii  occurs  on  surf  exposed  rocks 
surrounded  by  sandy  areas  in  the  intertidal  zone  and  the 
shallow  sublittoral  zones,  a habitat  entirely  unlike  that  of  the 
others.  I missed  seeing  this  species  because  I did  not  examine 
this  habitat  in  Chile  in  1975.  However,  the  occurrence  of  F. 
bridgesii  has  been  detailed  by  Bretos  ( 1 979).  Shells  are  usually 
overgrown  with  an  algal  mat.  Only  rarely  do  shells  have 
attachment  scars  of  Scurria,  indicating  that  they  usually  live 
deeper  than  the  mid-  to  upper  intertidal  zone  preferred  by 
Scurria.  The  Californian  acmaeid  limpet  Notoacmea  fenes- 
trata  (Reeve,  1855)  has  a comparable  habitat  on  rocks  sur- 
rounded by  sand. 

Distribution.  Isla  Guanape,  Peru  (8°32'  S)  (LACM  74-3, 
McLean),  to  Rio  Bio-bio,  Concepcion  Province,  Chile  (36°48' 
S)  (LACM  75-35,  McLean).  There  is  but  a single  record  from 


60  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


Figures  238  through  248.  Fissurel/a  bridgesii  Reeve,  1849.  Mature  shells.  (238)  3-5  m,  Isla  Guanape,  Peru.  LACM  74-3,  66.0  x 46.0  x 
15.0  mm.  (239)  Iquique,  Chile.  LACM  90802,  22.4  x 15.7  x 5.3  mm.  (240)  Same  locality.  LACM  90802,  20.0  x 12.8  x 5.9  mm.  (241) 
Iquique,  Chile.  LACM  90803,  65.5  x 41.3  x 14.5  mm.  (242)  Paposo,  Antofagasta  Province,  Chile.  LACM  54764,  44.4  x 28.6  x 9.3  mm. 
(243)  Quintero,  Valparaiso  Province,  Chile.  USNM  48221,  89.3  x 57.0  x 23.5  mm.  (244)  Paralectotype,  F.  bridgesii  Reeve.  Quintero,  Val- 
paraiso Province,  Chile.  BMNH  197566,  77.6  x 49.1  x 1 14.3  mm.  (245)  Lectotype,  F.  bridgesii  Reeve.  Quintero,  Valparaiso  Province,  Chile. 
BMNH  197566,  68.0  x 47.5  x 13.7  mm.  (246)  Valparaiso,  Chile.  USNM  56255,  33.0  x 19.5  x 10.2  mm.  (247)  Valparaiso,  Chile.  AMNH 
20055,  52.8  x 36.0  x 10.4  mm.  (248)  Rio  Bio-bio,  Concepcion  Province,  Chile.  LACM  75-35,  42.4  x 26.5  x 9.0  mm. 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  61 


249  253 


Figures  249  through  253.  Fissurella  bridgesii  Reeve,  1 849.  Living  and  preserved  specimens,  mantle  lobe  and  radula.  (249)  Three  living 
specimens  in  place.  Tocopilla,  Antofagasta  Province,  Chile,  photo  courtesy  A.  Viviani.  (250)  Mantle  lobe.  Iquique,  Chile.  LACM  90803,  length 
15  mm.  (251)  Mantle  lobe.  3-5  m,  Isla  Guanape,  Peru.  LACM  74-3,  length  20  mm.  (252)  Preserved  specimen.  Iquique,  Chile.  LACM  90803, 
shell  length  65.5  mm.  (253)  Air-dried  radula.  Iquique,  Chile.  LACM  90804,  width  3.5  mm,  shell  length  74.4  mm. 


Peru,  the  living  specimen  I collected  in  1974  at  Isla  Guanape 
(Fig.  238).  I have  examined  specimens  from  the  following 
localities  in  Chile:  Iquique,  Paposa,  Quintero,  Valparaiso, 
and  Rio  Bio-bio,  near  Concepcion. 

Number  of  Lots  Examined.  15  (LACM  8,  AMNH  1,MACN 
1,  MNHN  3,  USNM  2).  This  species  is  uncommon  in  the 
collections  examined. 

Taxonomic  History.  Accounts  of  F.  bridgesii  in  the  liter- 
ature prior  to  the  report  of  Bretos  (1979)  were  copies  of  the 
original  description  of  Reeve  (1849).  Dell  (1971)  considered 
this  taxon  a synonym  of  F.  latimarginata.  The  validity  of 
the  species  was  confirmed  in  1977,  when  I compared  a pre- 
served specimen  sent  to  me  for  identification  by  M.  Bretos 
with  the  syntypes  from  the  British  Museum  then  on  loan  at 
the  LACM.  Although  I was  unable  to  find  living  examples 


of  this  species  during  my  fieldwork  in  Chile,  puzzling  beach- 
worn  shells  were  collected  at  several  localities,  and  I later 
discovered  that  I had  purchased  live  specimens  at  the  market 
in  Iquique.  The  specimen  I had  earlier  collected  in  Peru  at 
Isla  Guanape  was  then  recognized  as  F.  bridgesii.  The  spe- 
cialized habitat  of  this  species  accounts  for  its  scarcity  in 
collections.  It  may  be,  however,  that  Ramirez-Boehme  (1974) 
had  recognized  the  species,  because  his  key  mentions  the 
characteristic  upwardly  beveled  margin  (“bordes  laterales  re- 
fiejados  hacia  arriba”),  a feature  not  discussed  elsewhere  in 
the  literature. 

Abundance  and  Use.  Fissurella  bridgesii  is  used  for  food 
at  Iquique,  and  it  may  have  more  economic  importance  than 
suggested  by  the  paucity  of  specimens  in  collections.  Ac- 
cording to  M.  Bretos,  it  is  known  to  the  fishermen  as  the 


62  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


“lapa  jerguilla,”  or  the  “lapa  de  arena”  (sand).  The  jerguilla 
is  a fish,  Aplodactylus  punctatus,  which  has  a color  pattern 
resembling  that  of  the  body  of  F.  bridgesii. 

Characteristics  and  Variability.  The  most  characteristic 
features  of  F.  bridgesii  are  the  lack  of  regularly  defined  radial 
sculpture  in  mature  shells,  a relatively  large  foramen,  a faintly 
rayed  pattern  on  a gray-brown  ground,  and  the  broad,  flat 
margin,  which  is  beveled  upward  in  young,  growing  speci- 
mens. Some  specimens  have  an  elongate  foramen  that  is 
bidentate  on  the  sides;  others  of  the  same  size  may  have  a 
more  oval  foramen.  Color  patterns  include  faintly  rayed  spec- 
imens and  some  uniformly  gray  shells  lacking  traces  of  rays. 
The  northernmost  specimen  from  Isla  Guanape,  Peru,  lacks 
rays  (Fig.  238). 

Affinity  and  Comparisons.  Fissurella  bridgesii  most  resem- 
bles F.  latimarginata.  The  normal  weakly  rayed  pattern  of 
F.  bridgesii  can  be  related  to  the  unusual  rayed  color  form 
of  F.  latimarginata.  Both  have  a broad  margin  and  a lighter 
outer  zone  to  the  calcitic  layer,  but  only  F.  bridgesii  has  an 
upwardly  beveled  margin.  Both  species  have  similar  elabo- 
ration of  the  tongue-shaped  papillae  of  the  mantle  edge,  more 
so  than  do  other  species.  In  its  light  outer  shell  layer  it  also 
resembles  F.  limbata,  but  may  be  distinguished  from  that 
species  in  lacking  the  purple  coloration  to  the  shell.  Fissurella 
bridgesii  also  resembles  F.  pulchra  in  its  low  profile,  weakly 
contrasting  pattern  of  rays,  and  the  lighter  outermost  zone 
to  the  calcitic  layer.  It  differs  in  lacking  the  speckled  pattern 
of  F.  pulchra  and  in  having  a larger  foramen  and  a more 
pronounced  upward-beveled  margin.  Although  the  size  of 
the  foramen  varies  somewhat  in  F.  bridgesii,  it  is  always 
larger  than  that  of  similarly  sized  specimens  of  F.  pulchra. 
Before  I understood  F.  bridgesii,  my  guess  was  that  young 
beach-worn  specimens,  such  as  the  one  from  Concepcion 
(Fig.  248),  were  most  likely  to  be  variants  of  F.  pulchra. 

Synonymy  and  Types.  There  are  four  syntype  specimens 
of  F.  bridgesii  Reeve,  1 849,  from  Quintero,  Valparaiso  Prov- 
ince, Chile  (32°47'  S),  BMNH  197566,  lengths  77.7,  68.0, 
61.8,  and  46.0  mm.  The  68.0  mm  specimen  was  figured  by 
Reeve  (1849),  Sowerby  II  (1862),  and  Dell  (1971),  and  is 
here  designated  the  lectotype  (Fig.  245);  the  larger  77.7  mm 
specimen  has  been  figured  only  by  Sowerby  II  (1862).  These 
two  large  specimens  (Figs.  244,  245)  illustrate  two  of  the 
most  variable  features  of  the  species.  They  are  nearly  of  the 
same  width,  but  one  is  much  more  elongate.  The  larger, 
elongate  specimen  has  an  oval  foramen;  the  broader  speci- 
men has  a bidentate  foramen.  The  two  remaining  paralec- 
totypes  are  broad;  the  61.8  mm  specimen  has  an  oval  fora- 
men; the  46.0  mm  specimen  has  an  elongate  foramen  with 
the  bidentate  structure  nearly  imperceptible.  Shape  of  the 
foramen  is  therefore  not  correlated  with  shell  proportions. 

Fissurella  pulchra  Sowerby,  1835 

Figures  254-267 

Fissurella  pulchra  Sowerby,  1835a:  124;  Sowerby,  1835b:3, 
fig.  24;  Reeve,  1849,  pi.  2,  fig.  9;  Hupe,  1854:244;  Philippi, 
1860:181;  Sowerby  II,  1862:184,  fig.  31;  Pilsbry,  1890: 
151,  pi.  33,  fig.  50;  Dali,  1909:242;  Riveros-Zuniga,  1951: 


120,  fig.  27;  Dell,  1971:191,  pi.  3,  fig.  5;  Ramirez-Boehme, 

1974:30  [key]. 

Shell.  Medium-sized  (35-75  mm  mature  length),  consis- 
tently low;  outline  elongate  oval,  tapered  anteriorly,  sides 
slightly  raised  relative  to  ends.  Radial  sculpture  in  early  growth 
stages  consisting  of  low,  rounded  primary  ribs,  becoming 
obsolete  when  shell  reaches  length  of  20  mm;  mature  shell 
smooth  except  for  faint  radial  striae.  Color  grayish  lavender 
to  pink,  with  alternating  lighter  and  darker  rays;  entire  surface 
with  fine  reddish  speckles  or  zigzag  markings  especially  pro- 
nounced near  foramen;  concentric  interruptions  to  rays 
changing  color  from  pink  to  gray  or  brown  in  some.  Margin 
of  moderate  width,  flat,  reddish  gray,  zoned  to  make  edge 
slightly  lighter  in  color;  broken  shells  showing  lighter  outer 
zone  of  calcitic  layer;  margin  becoming  very  narrow  in  large, 
full  grown  shells.  Foramen  elongate  and  tripartite  in  young 
shells,  only  slightly  less  elongate  in  mature  shells;  interior 
callus  bordered  by  pink  colored  ring  in  attachment  region. 

Juvenile  Shell.  Primary  ribs  rounded,  coinciding  with 
lighter  rays,  speckled  pattern  conspicuous.  Sides  of  shell  raised 
bordering  foramen,  indicating  that  earliest  stage  is  more  con- 
ical. 

Mantle  and  Foot.  Not  retractable  in  flattened  shell;  mantle 
lobe  normally  extending  well  over  shell  edge.  Cephalic  ten- 
tacles lavender,  yellowish  at  tips.  Mantle  lobe  pinkish  gray 
or  brown,  faintly  banded  to  match  rayed  pattern,  lined  con- 
centrically with  brown.  Papillae  of  upper  edge  well  devel- 
oped, those  of  lower  edge  more  numerous  and  smaller.  Side 
of  foot  same  color,  tubercles  well  developed. 

Habitat.  Low  intertidal  and  sublittoral  zones  in  crevices 
or  on  undersides  of  large  rocks  in  deep  tidepools,  protected 
from  direct  exposure  to  surf.  Shells  are  free  of  algal  incrus- 
tations; epibiotic  organisms  on  the  shell  are  bryozoans  and 
spirorbid  polychaetes. 

Distribution.  Salaverry,  Peru  (8°14'  S)  (USNM  368490,  W. 
Schmitt),  to  Rio  Bio-bio,  Concepcion  Province,  Chile  (36°48' 
S)  (LACM  75-35,  McLean).  Except  for  F.  bridgesii,  this  is 
the  least  common  species  in  the  Peruvian  Faunal  Province. 
I have  found  small  specimens  at  most  localities  in  central 
Peru.  In  northern  Chile  it  was  rare  at  Iquique;  at  Antofagasta 
there  was  no  trace  of  it,  even  in  the  beach-worn  shell  debris. 
It  was  more  common  in  central  Chile,  where  I found  living 
specimens  on  undersides  of  boulders  in  tidepools  at  Los 
Molles,  Aconcagua  Province.  Beach-worn  specimens  were 
found  at  other  localities  in  central  Chile. 

Number  of  Lots  Examined.  21  (LACM  13,  AMNH  1,ANSP 
3,  USNM  4,  none  at  MACN  or  MNHN). 

Taxonomic  History.  Most  authors  have  merely  copied  the 
original  account  of  F.  pulchra,  except  for  Pilsbry  ( 1 890),  who 
emphasized  the  characteristic  speckled  pattern.  The  habitat 
is  cryptic,  and  beach-worn  shells  are  sufficiently  scarce  that 
the  species  has  escaped  notice  in  recent  years.  Riveros-Zuniga 
(1951)  merely  quoted  previous  authors,  and  Pena  (1970)  did 
not  mention  it.  The  collection  from  Iquique  of  Marincovich 
(1973)  did  not  include  it.  Ramirez-Boehme  ( 1974)  included 
it  in  his  key  but  did  not  mention  the  conspicuous  speckles. 

Abundance  and  Use.  The  species  is  sufficiently  uncommon 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  63 


Figures  254  through  259.  Fissurella  pulchra  Sowerby,  1835.  Mature  shells.  (254)  Miraflores,  Lima  Province,  Peru.  LACM  71-187,  24.2  x 
13.5  x 4.  | mm.  (255)  Pupudo,  Aconcagua  Province,  Chile.  LACM  54656,  78.5  x 56.0  x 17.0  mm.  (256)  Paralectotype,  F.  pulchra  Sowerby. 
Valparaiso,  Chile.  BMNH  197580,  39.6  x 25.8  x 6.6  mm.  (257)  Lectotype.  F.  pulchra  Sowerby.  Valparaiso,  Chile.  BMNH  197580,  62.2  x 
40.2  x 12.8  mm.  (258)  Cartagena,  Santiago  Province,  Chile.  LACM  75-34,  68.5  x 50.8  x 14.3  mm.  (259)  Rio  Bio-bio,  Concepcion  Province, 
Chile.  LACM  75-35,  37.5  x 23.3  x 6.6  mm. 


to  be  negligible  as  a food  resource.  Large  specimens  are  par- 
ticularly uncommon;  living  specimens  that  I found  were  about 
half  the  size  of  shells  in  the  type  lot,  although  I found  some 
beach-worn  shells  that  approached  the  maximum  size.  The 
largest  specimen  1 have  examined  is  78.5  mm  in  length,  from 
Papudo,  Aconcagua  Province,  Chile,  donated  to  the  LACM 
by  J.  Ramirez-Boehme  (Fig.  255).  The  species  has  no  com- 
mon name  in  northern  Chile,  according  to  M.  Bretos. 

Characteristics  and  Variability.  Fissurella  pulchra  is  aptly 
named.  It  is  rayed  in  pinkish  brown  and  gray  and  is  the  only 
species  in  which  a speckled  pattern  persists  through  all  growth 
stages.  It  is  one  of  the  least  variable  species;  I have  noticed 
no  unusual  color  forms.  Shells  are  consistently  low  and  ta- 
pered anteriorly.  Most  shells  are  relatively  smooth,  although 
the  weak  primary  ribs  may  be  more  pronounced  in  some 
specimens. 

The  radula  of  F.  pulchra  (Figs.  266,  267)  is  the  most  dis- 
tinctive among  the  Peruvian-Magellanic  species.  The  inner 
laterals  have  longer  overhanging  cusps  than  the  other  species. 


and  the  enlarged  outer  lateral  is  unique  in  having  a concave 
edge  to  the  second  cusp. 

Affinity  and  Comparisons.  On  shell  characters,  F.  pulchra 
surely  hts  within  the  group  having  broad  primary  ribs  and 
no  secondary  ribs.  It  is  closest  to  F.  bridgesii,  with  which  it 
shares  similar  proportions,  general  overall  coloration,  and 
faint  rays.  It  differs  in  its  speckled  pattern,  more  consistently 
narrowed  foramen,  and  pink  bordered  internal  callus.  The 
margin  is  less  broad  than  that  of  F.  bridgesii;  it  is  similarly 
beveled  upward  but  to  a lesser  extent  than  in  F.  bridgesii. 
The  tapered  anterior  end  of  F.  pulchra  resembles  the  con- 
dition of  the  otherwise  not  closely  related  F.  latimarginata 
and  F.  cumingi. 

Synonymy  and  Types.  There  are  5 syntypes  of  F.  pulchra 
Sowerby,  1835,  from  Valparaiso,  Chile,  BMNH  197580, 
lengths  70.8  (broken  in  half),  62.2,  48.7,  39.6,  and  24.0  mm. 
The  largest  of  these  has  not  been  figured.  The  62.2  mm 
specimen  was  figured  originally  by  Sowerby  (1835b),  then  by 
Reeve  (1849)  and  by  Dell  (1971);  it  is  here  designated  the 


64  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


Figures  260  through  267.  Fissurella  pulchra  Sowerby,  1835.  Living  specimens,  mantle  lobe,  juvenile  shells,  radulae.  (260)  Living  specimen 
with  mantle  expanded,  in  tidepool.  Cumbres  Borascosas,  Tarapaca  Province,  Chile.  LACM  75-14.  (261)  Same  specimen,  ventral  view.  (262) 
Living  specimen  on  underside  of  overturned  boulder.  Los  Molles,  Aconcagua  Province,  Chile.  LACM  75-28.  (263)  Mantle  lobe.  Cumbres 
Borascosas,  Tarapaca  Province,  Chile.  LACM  75-14,  8 mm.  (264)  Juvenile  specimen.  Los  Molles,  Tarapaca  Province,  Chile.  LACM  75-28, 
5.8  x 3.5  x 1.3  mm.  (265)  Juvenile  shell.  Pucusana,  Lima  Province,  Peru.  LACM  72-76,  11.7  x 6.7  x 2.0  mm.  (266)  Radula  of  small  shell. 
Same  locality.  LACM  72-76,  1.5  mm,  shell  length,  22.1  mm.  (267)  SEM  view  of  radula.  Los  Molles,  Aconcagua  Province,  Chile.  LACM  75- 
28,  width  of  field  0.6  mm. 


lectotype.  The  lectotype  and  the  39.6  mm  paralectotype  are 
figured  here  (Figs.  256,  257). 

LACM  LOCALITIES  FOR  FIGURED 
SPECIMENS 

[All  collections  from  intertidal  zone,  unless  otherwise  indi- 
cated.] 

AHF  375-35.  Bahia  Independencia,  lea  Province,  Peru(  14°14' 
S,  76°12.7'  W),  13  January  1935. 

AHF  380-35.  Bahia  Independencia,  lea  Province,  Peru  (14°  14' 
S,  76°08.5'  W),  14  January  1935. 

AHF  828-38.  Bahia  San  Juan,  lea  Province,  Peru  (15°20.7' 
S,  75°09.3'  W),  8 February  1938. 


62-26.  Ancud,  Chiloe  Province,  Chile  (41°52'  S,  73°05'  W), 
H.C.  McMillin,  24  March  1962. 

64-16.  Iquique  (near  Ave.  Baquedena),  Tarapaca  Province, 
Chile  (20°13'  S,  70°10'  W),  L.  Marincovich,  June  through 
August,  1964. 

66-46.  Renaca,  Vina  del  Mar,  Valparaiso  Province,  Chile 
(33°06'  S,  7 1°50'  W),  R.  Seapy,  24  July  1966. 

70- 68.  Iquique  (10  km  S),  Tarapaca  Province,  Chile  (20°15' 
S,  70°09'  W),  L.  Marincovich,  July,  1970. 

71- 187.  Miraflores,  Lima  Province,  Peru  (12°08'  S,  77°04.5' 
W),  T.  Bratcher,  24  February  1971. 

7 1 -277.  Bahia  York,  Isla  de  los  Estados,  Argentina  (54°47. 1 1 ' 
S,  64°1 7.9'  W),  5 May  1971. 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  65 


7 1 -284.  Puerto  San  Juan  del  Salvamento,  Isla  de  los  Estados, 
Argentina  (54°43.9'  S,  63°52'  W),  13  May  1971. 

72-76.  0-5  m,  Pucusana,  Lima  Province,  Peru  (12°30'  S, 
76°49'  W),  J.H.  McLean,  30  March  1972. 

72-77.  Laguna  Granda,  lea  Province,  Peru  ( 1 4°  1 8'  S,  76°15' 
W),  J.H.  McLean,  31  March  1972  (beach-worn  shells). 
72-79.  Paracas,  lea  Province,  Peru  (13°49'  S,  76°14.5'  W). 
J.H.  McLean,  2 April  1972  (beach-worn  shells). 

74-2.  NE  side,  Isla  Guanape,  La  Libertad  Province,  Peru 
(08°32'  S,  78°58'  W),  J.H.  McLean,  18  January  1974. 

74-3.  3-5  m,  NE  side,  Isla  Guanape,  La  Libertad  Province, 
Peru  (08°32'  S,  78°58'  W),  J.H.  McLean,  18  January  1974. 
74-21.  1-2  m,  Playa  Hermosa,  Ancon,  Lima  Province,  Peru 
(1 1°47'  S,  77°1 1.5'  W),  J.H.  McLean,  26  January  1974. 

74- 24.  \-4  m,  Isla  San  Lorenzo,  Lima  Province,  Peru(12°06.7' 
S,  77°13'  W),  J.H.  McLean,  29  January  1974. 

75- 10.  Pozo  Toyo  (S  of  Iquique),  Tarapaca  Province,  Chile 
(20°25'  S,  70°  10.5'  W),  J.H.  McLean,  29  September  and  1 
October  1975. 

75-12.  Iquique  (at  Marine  Laboratory,  Universidad  del 
Norte),  Tarapaca  Province,  Chile  (20°1 5.5'  S,  70°08'  W),  J.H. 
McLean,  2 October  1975. 

75-14.  Cumbres  Borascosas,  Tarapaca  Province,  Chile 
(20°42'S,  70°1  1.5'  W),  J.H.  McLean,  3 October  1975. 

75-15.  Antofagasta  (S  end  of  city),  Antofagasta  Province, 
Chile  (23°42'  S,  70°27'  W),  J.H.  McLean,  5 and  6 October 
1975. 

75-16.  Bahia  Moreno,  Antofagasta  Province,  Chile  (23°28' 
N,  70°31'  W),  J.H.  McLean,  7 October  1975  (beach  shells). 
75-17.  [Shore  opposite]  Isla  Santa  Maria,  Antofagasta  Prov- 
ince, Chile  (23°25'  S,  70°36'  W). 

75-18.  Punta  Jara,  Antofagasta  Province,  Chile  (23°49'  S, 
70°29'  W),  J.  Tomicic,  8 October  1975. 

75-19.  Los  Colorados,  Antofagasta  Province,  Chile  (23°29' 
N,  70°22'  W),  J.H.  McLean,  9 October  1975. 

75-20.  2-5  m,  Antofagasta  (S  end  of  city),  Antofagasta  Prov- 
ince, Chile  (23°42'  S,  70°27'  W),  J.H.  McLean,  10  October 
1975. 

75-21.  7-20  m,  El  Rincon  de  Mejillones,  Antofagasta  Prov- 
ince, Chile  (23°02'  S,  70°31'  W),  J.H.  McLean,  11  October 
1975 

75-23.  2-4  m,  El  Rincon  de  Mejillones,  Antofagasta  Prov- 
ince, Chile  (23°05' S,  70°30'  W),  J.  Tomicic,  12  October  1975. 
75-25.  Bahia  Herradura,  Coquimbo  Province,  Chile  (29°59' 
S,  7 1°22'  W),  J.H.  McLean,  14  October  1975. 

75-27.  Bahia  El  Teniente,  Coquimbo  Province,  Chile  (30°58' 
S,  7 1°39'  W),  J.H.  McLean,  15  October  1975. 

75-28.  Los  Molles,  Aconcagua  Province,  Chile  (32°14'  S, 
71°32'  W),  J.H.  McLean,  16  through  18  October  1975. 
75-29.  Los  Modes,  Aconcagua  Province,  Chile  (32°14'  S, 
71°32'  W),  J.H.  McLean,  16  October  1975  (shed  pile). 
75-30.  Montemar  (at  Estacion  de  Biologia  Marina),  Valpa- 
raiso Province,  Chile  (32°57'  S,  71°32'  W),  J.H.  McLean,  19 
and  20  October  1975. 

75-31.  Islota  Concon,  N of  Vina  del  Mar,  Valparaiso  Prov- 
ince, Chile  (32°52'  S,  71°33'  W),  J.H.  McLean,  21  and  22 
October  1975. 


75-34.  Cartagena,  Santiago  Province,  Chile  (33°33'  S,  71°38' 
W),  J.H.  McLean,  23  October  1975. 

75-35.  Rio  Bio-bio,  Concepcion  Province,  Chile  (36°48'  S, 
73°1 1 ' W),  J.H.  McLean,  29  October  1975. 

75-36.  Mehuin,  Valdivia  Province,  Chile  (39°23'  S,  73°14' 
W),  J.H.  McLean,  31  October  and  2 November  1975. 
75-37.  [Island  off]  Mehuin,  Valdivia  Province,  Chile  (39°26' 
S,  73°16'  W),  J.H.  McLean,  1 November  1975. 

75-39.  Pargua,  Canal  de  Chacao,  Llanquihue  Province,  Chile 
(41°47'  S,  73°28'  W),  J.H.  McLean,  3 November  1975. 
75-40.  Guabun,  Isla  de  Chiloe,  Chiloe  Province,  Chile  (41°50' 
S,  74°02'  W),  J.H.  McLean,  4 November  1975. 

75-41.  Pumalin,  Golfo  Corcovado,  Chiloe  Province,  Chile 
(42°42'  S,  72°52'  W),  J.H.  McLean,  4 through  6 November 
1975. 

75-42.  2-4  m,  Punta  Estero,  Isla  Talcon,  Golfo  Corcovado, 
Chiloe  Province,  Chde  (42°46'  S,  72°56'  W),  J.H.  McLean, 
6 November  1975. 

75-43.  4-13  m,  Islota  Nihuel,  Golfo  Corcovado,  Chiloe 
Province,  Chile  (42°38'  S,  72°57'  W),  J.H.  McLean,  7 No- 
vember 1975. 

75-44.  Islota  Nihuel,  Golfo  Corcovado,  Chiloe  Province,  Chile 
(42°38'  S,  72°57'  W),  J.H.  McLean,  7 November  1975. 
75-45.  Quellon,  Isla  de  Chiloe,  Chiloe  Province,  Chile  (43°09' 
S,  73°37'  W),  J.H.  McLean,  8 November  1975. 

75-47.  Isla  Laitec,  off  SE  side  Isla  de  Chiloe,  Chiloe  Province, 
Chile  (43°14'  S,  73°36'  W),  9 November  1975. 

75-48.  Fuerte  Bulnes,  Peninsula  Brunswick,  Magellan  Strait, 
Magallanes  Province,  Chile  (53°38'  S,  70°54.5'  W),  J.H. 
McLean,  16  November  1975. 

75-49.  Puerto  el  Hambre,  Peninsula  Brunswick,  Magellan 
Strait,  Magallanes  Province,  Chile  (53°37'  S,  70°56'  W),  J.H. 
McLean,  16  and  19  November  1975. 

78-88.  Punta  Ninfas,  Golfo  Nuevo,  Chubut  Province,  Ar- 
gentina (42°56. 5' S,  64°19.5' W),  J.H.  McLean,  18  July  1978. 
78-90.  Punta  Cracker,  Golfo  Nuevo,  Chubut  Province,  Ar- 
gentina (42°56.5'  S,  64°30'  W),  J.H.  McLean,  19  July  1978. 

ACKNOWLEDGMENTS 

I am  especially  indebted  to  those  who  made  the  arrangements 
for  fieldwork  in  South  America. 

In  Peru,  I was  aided  by  Enrique  M.  del  Solar  of  Lima,  and 
Norma  Chirichigno  and  Victor  Alamo,  of  the  Instituto  del 
Mar  in  Callao.  Collecting  excursions  were  arranged  out  of 
Lima  in  March,  1972,  and  in  January,  1974, 1 was  privileged 
to  join  a cruise  of  the  Peruvian  Government  research  vessel 
SNP-1  to  Isla  Guanape  and  the  Islas  Lobos  Afueras.  Other 
Peruvian  biologists  who  have  been  helpful  through  exchanges 
and  correspondence  include  Wenceslao  Medina,  G.  Mario 
Pena,  and  Carlos  Paredes  Quiroz. 

In  Chile,  I joined  the  Marine  Communities  Expedition  of 
Robert  T.  Paine  of  the  University  of  Washington,  October- 
November,  1975.  Expenses  were  partially  underwritten  by 
the  National  Science  Foundation  (DES  75-14378,  R.T.  Paine, 
principal  investigator).  Other  members  of  the  expedition, 
Carol  Sturgess,  Ken  Sebens,  and  Tom  Suchanek,  all  from  the 
University  of  Washington,  also  helped  in  the  collecting. 


66  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


Arrangements  in  Chile  were  made  by  Juan  Carlos  Castilla, 
Universidad  Catolica  de  Chile,  Santiago,  who  joined  us  for 
part  of  the  expedition,  made  the  facilities  of  the  Universidad 
Catolica  available  to  us,  and  arranged  to  ship  the  collections 
to  me.  Others  who  assisted  us  were  C.A.  Viviani  and  Marta 
Bretos,  then  of  Universidad  del  Norte,  Iquique;  Jorge  To- 
rnicic,  Universidad  de  Antofagasta,  Antofagasta;  Miguel  Pa- 
dilla and  Luis  Ramorino,  Instituto  de  Oceanologia,  Univer- 
sidad de  Valparaiso,  Valparaiso;  Olga  Aracena,  Universidad 
de  Concepcion,  Concepcion;  Hugo  Campos,  Universidad 
Austral  de  Chile,  Valdivia;  and  Italo  Compodonico,  Instituto 
de  la  Patagonia,  Punta  Arenas.  Every  effort  was  made  to  assist 
us  by  our  Chilean  hosts. 

Fieldwork  in  Argentina  in  July,  1978,  was  arranged  by 
William  J.  Zinsmeister  of  Ohio  State  University,  chief  sci- 
entist on  cruise  783  of  the  R/V  HERO  to  the  vicinity  of  the 
Peninsula  Valdez. 

Collections  made  by  Luie  Marincovich  in  Iquique  in  1964 
and  1980  were  instrumental  in  introducing  me  to  the  Pe- 
ruvian-Magellanic  fauna. 

I also  thank  Paul  Dayton  of  Scripps  Institution  of  Ocean- 
ography for  making  a very  productive  effort  at  my  request 
to  collect  mollusks  during  his  research  cruises  in  southern 
Chile  on  the  R/V  HERO  in  1972  and  1973. 

Edmundo  Martinez,  of  Antofagasta,  kindly  provided  me 
with  fossil  specimens  of  Fissurella  species  from  that  region 
of  Chile. 

I am  particularly  indebted  to  K.M.  Way  of  the  British 
Museum  for  arranging  the  loan  of  the  types  of  Fissurella 
preserved  in  that  institution  and  providing  much  necessary 
information  through  correspondence. 

Philippe  Bouchet  and  B.  Metivier  of  the  Paris  Museum 
kindly  searched  for  the  types  and  provided  loan  material  of 
a number  of  other  species  described  by  early  French  authors. 

In  1978  I visited  the  Museo  Nacional  de  Historia  Natural, 
Santiago,  where  I was  assisted  by  Maria  Codoceo  and  Ni- 
baldo  Bahamonde.  On  the  same  trip  I visited  the  Museo 
Argentino  de  Ciencias  Naturales,  Buenos  Aires,  where  I was 
assisted  by  Martinez  Fontes. 

I thank  William  K.  Emerson  of  the  American  Museum  of 
Natural  History,  Joseph  Rosewater  and  Richard  S.  Houbrick 
of  the  United  States  National  Museum,  and  Robert  Robert- 
son and  George  M.  Davis  of  Academy  of  Natural  Sciences, 
Philadelphia,  for  making  the  collections  of  their  institutions 
available  to  me  and  arranging  loans  of  specimens.  Loans  were 
also  arranged  by  Kenneth  J.  Boss  of  the  Museum  of  Com- 
parative Zoology,  Harvard,  and  Richard  K.  Dell  of  the  Na- 
tional Museum  of  New  Zealand. 

Marta  Bretos,  now  of  the  Universidad  de  la  Frontera,  Te- 
muco,  Chile,  has  reviewed  the  manuscript,  sent  specimens, 
and  supplied  me  with  the  common  names  of  Fissurella  species 
as  used  in  northern  Chile. 

Polychaete  shell  borers  were  identified  by  K.  Fauchald  of 
the  U.S.  National  Museum  of  Natural  History. 

C.J.  Risso-Dominguez  of  Buenos  Aires  provided  me  with 
specimens  and  information  about  the  occurrence  of  Fissu- 
rella species  in  Argentina  and  reviewed  an  early  draft  of  the 
manuscript. 


Carole  S.  Hickman,  University  of  California,  Berkeley, 
provided  the  SEM  micrographs  of  radulae;  Heinz  A.  Low- 
enstam,  California  Institute  of  Technology,  Pasadena,  con- 
tributed the  SEM  micrographs  of  the  shell  of  a small  speci- 
men of  Fissurella  latimarginata  (Figs.  4-7).  Edward  Wilson 
of  the  LACM  section  of  Invertebrate  Paleontology  cut  spec- 
imens for  the  examination  of  shell  layers. 

Photographs  of  shells  and  air-dried  radulae  are  the  work 
of  Bertram  C.  Draper,  museum  volunteer.  Museum  photog- 
rapher Dick  Meier  photographed  the  cut  shells.  Museum 
illustrators  Mary  Butler  and  Caryl  Maloof  assisted  in  the 
preparation  of  the  figures.  Museum  volunteer  Jo-Carol  Ram- 
saran  helped  in  curatorial  tasks  and  library  searches  necessary 
to  the  work. 

I thank  H.  Andrade  for  translating  the  abstract  to  Spanish 
and  Fernando  Jara  for  reviewing  the  ecological  remarks. 

Comments  and  criticisms  of  various  drafts  of  this  paper 
have  been  received  from  Eugene  Coan,  Cliff  Coney,  J.  Chris- 
tiaens,  Myra  Keen,  Patrick  I.  LaFollette,  David  R.  Lindberg, 
and  Robert  T.  Paine.  The  submitted  manuscript  was  re- 
viewed by  William  K.  Emerson,  Barry  Roth,  and  Edward 
Wilson. 

LITERATURE  CITED 

Acuna,  E.  1977.  Estudio  preliminar  de  edad  y crecimiento 
de  Fissurella  latimarginata  (Sowerby,  1834)  en  Toco- 
pilla,  Chile  (Mollusca,  Gastropoda,  Fissurellidae).  Re- 
vista Biologia  Marina,  Departamento  de  Oceanologia, 
Universidad  de  Chile,  Valparaiso  1 6(2):  1 17-124. 
Boggild,  O.B.  1930.  The  shell  structure  of  the  mollusks. 
Det  Kongelige  Danske  Videnskabernes  Selskabs  Skrif- 
ter,  Niende  Raekke,  Naturvidenskabelig  og  Mathema- 
tisk  Afdeling  9,  Raekke  2:231-326,  pis.  1-15. 

Boutan,  L.  1885.  Recherches  sur  l’anatomie  et  le  devel- 
oppement  de  la  Fissurella.  Archives  de  Zoologie  Expe- 
rimental et  Generale,  series  2,  tome  3,  memoire  4:1- 
173,  pis.  31-44. 

Bretos,  M.  1978.  Growth  in  the  keyhole  limpet  Fissurella 
crassa  Lamarck  (Mollusca:  Archaeogastropoda)  in 
northern  Chile.  The  Veliger  21(2):268-273. 

. 1979.  Observaciones  sobre  Fissurella  bridgesii 

Reeve,  1849,  en  Tarapaca,  Norte  de  Chile.  Ciencia  y 
Tecnica  del  Mar,  Cona  4:53-60. 

. 1980.  Age  determination  in  the  keyhole  limpet 

Fissurella  crassa  Lamarck  (Archaeogastropoda:  Fissu- 
rellidae), based  on  shell  growth  rings.  Biological  Bulletin 
159:606-612. 

. 1982.  Biologia  de  Fissurella  maxima  Sowerby 

(Mollusca:  Archaeogastropoda)  en  el  Norte  de  Chile.  1. 
Caracteres  generales,  edad  y crecimiento.  Cahiers  de 
Biologie  Marine  23: 1 59- 1 70. 

. 1983.  The  biology  of  Fissurella  maxima  Sowerby 

(Mollusca:  Archaeogastropoda)  in  northern  Chile.  2. 
Notes  on  its  reproduction.  Biological  Bulletin  1 65:559— 
568. 

Bretos,  M.,  and  C.  Jiron.  1980.  Trematodes  in  Chilean 
fissurellid  molluscs.  The  Veliger  22(3):293. 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  67 


Bruguiere,  J.  1789.  Histoire  naturelle  des  vers.  Paris,  v.  1, 
part  1,  344  pp. 

Carcelles,  A.R.  1950.  Catalogo  de  los  moluscos  marinos 
de  la  Patagonia.  Anales  del  Museo  Nahuel  Huapi  2:41  — 
92,  pis.  1-6. 

. 1953.  Catalogo  de  la  malacofauna  Antartica  Ar- 
gentina. Anales  del  Museo  Nahuel  Huapi  3:1 50-250,  pis. 
1-5. 

Carcelles,  A.R. , and  S. I.  Williamson.  1951.  Catalogo  de  los 
moluscos  marinos  de  la  provincia  Magallanica.  Revista 
del  Instituto  Nacional  de  lnvestigacion  de  la  Ciencias 
Naturales,  Museo  Argentino  de  Ciencias  Naturales 
" Bernardino  Rivadavia,  ” Ciencias  Zoologicas  2(5):225- 
283. 

Castilla,  J.C.  1981.  Perspectivas  de  investigacion  en  es- 
tructura  y dinamica  de  comunidades  intermareales  ro- 
cosas  de  Chile  Central.  II.  Depredadores  de  alto  nivel 
trofico.  Medio  Ambiente  5(1-2):  190-2 15. 

Castilla,  J.C.,  and  I.  Bahamondes.  1979.  Observaciones 
conducuales  y ecologicas  sobre  Lutra  felina  (Molina) 
1782  (Carnivora:  Mustelidae)  en  las  zonas  Central  y 
Centro-Norte  de  Chile.  Archivos  de  Biologla  y Medicina 
Experimental  1 1 2: 1 19-132. 

Christiaens,  J.  1973.  Les  Fissurelles  Europeennes— II.  Le 
genre  Fissurella  Bruguiere,  1789.  Informations  de  la  So- 
ciete  Beige  de  Malacologie,  series  2,  no.  5,  pp.  67-98. 

Dall.W.H.  1909.  Report  on  a collection  of  shells  from  Peru, 
with  a summary  of  the  littoral  marine  Mollusca  of  the 
Peruvian  zoological  province.  Proceedings  of  the  United 
States  National  Museum  37:147-294. 

Dance,  S.P.  1966.  Shell  collecting:  An  illustrated  history. 
Berkeley:  University  of  California  Press,  344  pp. 

Davila,  P.F.  1767.  Catalogue  systematique  et  raisonne  des 
curiosites  de  la  nature  et  de  l 'art.  qui  composent  la  cab- 
inet de  M.  Davila.  Paris,  3 vols. 

Dayton,  P.K.,  R.J.  Rosenthal,  L.C.  Mahen,  and  T.  Antezana. 
1977.  Population  structure  and  foraging  biology  of  the 
predaceous  Chilean  asteroid  Meyenaster  gelatinosus  and 
the  escape  biology  of  its  prey.  Marine  Biology  39:36 1 — 
370. 

Delessert,  B.  1841.  Recueil  de  coquilles  decrites  par  La- 
marck dans  son  Histoire  naturelle  des  animaux  sans  ver- 
tebras et  non  encore  figurees.  Paris,  40  pis.  (with  unnum- 
bered explanations). 

DelfR.K.  1971.  The  marine  Mollusca  of  the  Royal  Society 
Expedition  to  Southern  Chile,  1958-59.  Records  of  the 
Dominion  Museum  (Wellington)  7(1 7):  155-233. 

Deshayes,  G.P.  1830-1832.  Encyclopedic  Methodique. 
Histoire  naturelle  des  vers.  Paris,  v.  2,  594  pp. 

Dwyer,  K.R.,  and  D.R.  Lindberg.  1981.  The  topography, 
formation,  and  role  of  the  home  depression  of  the  limpet 
Collisella  scabra  Gould.  13th  Annual  Report,  Western 
Society  of  Malacologists,  p.  1 1 . 

Fretter,  V.,  and  A.  Graham.  1962.  British prosobranch  mol- 
luscs. London:  Ray  Society,  xvi  + 755  pp. 

. 1964.  Reproduction.  In  Physiology’  of  Mollusca. 

ed.  K.M.  Wilbur  and  C.M.  Yonge,  127-164.  New  York: 
Academic  Press. 


. 1976.  The  prosobranch  molluscs  of  Britain  and 

Denmark.  Part  1.  Pleurotomariacea,  Fissurellacea,  and 
Patellacea.  Journal  of  Molluscan  Studies.  Supplement  1 , 
1-37. 

Gallardo,  C.,  and  C.  Osorio.  1978.  Hiatella  solida  (Sow- 
erby,  1834)  (Mollusca:  Hiatellidae)  on  Concholepas con- 
cholepas  (Bruguiere,  1 789)  and  other  substrates.  The  Ve- 
liger  20(3):274-278. 

Gibson,  P.H.  1978.  Systematics  of  Dodecaceria  (Annelida: 
Polychaeta)  and  its  relation  to  the  reproduction  of  its 
species.  Zoological  Journal  of  the  Linnean  Society  63: 
275-287. 

Gmelin,  J.F.  1791.  Caroli  a Linne  systema  naturae  per 
regna  tria  naturae.  Editio  decima  tertia.  Leipzig,  v.  1, 
part  6,  Vermes,  pp.  3021-3910. 

Gould,  A.  A.  1846.  Descriptions  of  the  shells  collected  by 
the  United  States  Exploring  Expedition.  Proceedings  of 
the  Boston  Society  of  Natural  History  2:148-152. 

. 1852-1856.  Mollusca  and  shells.  United  States  Ex- 
ploring Expedition.  Boston,  v.  12,  509  pp.  [1852];  Atlas, 
16  pp.,  52  pis.  [1856], 

Gray,  J.E.  1839.  Molluscous  animals  and  their  shells.  In 
F.  W.  Beechey,  The  zoology  of  Capt.  Beechey’s  voyage 
...  to  the  Pacific  and  Behring’s  Straits  in  his  Majesty’s 
ship  “Blossom.  ” i-xii,  103-155,  pis.  33-44.  London. 

Herm,  D.  1969.  Marines  Pliozan  und  Pleistozan  in  Nord- 
und  Mittel-Chile  unter  besonderer  Beriicksichtigung  der 
Entwicklung  der  Mollusken-Faunen.  Zitteliana  2:1-158. 

Hickman,  C.S.  1981.  Evolution  and  function  of  asymmetry 
in  the  archaeogastropod  radula.  The  Veliger  23(3):  1 89— 
194,  2 pis. 

. 1984.  Implications  of  radular  tooth-row  functional 

integration  for  archaeogastropod  systematics.  Malaco- 
logia  25(1):  143-1 60. 

Hupe,  L.H.  1854.  Fauna  Chilena,  Moluscos.  In  Historia 
fisica  y politico  de  Chile,  ed.  C.  Gay.  Zoologia,  v.  8. 
Santiago,  499  pp.,  atlas,  v.  2,  pis.  1-8. 

Ihering,  H.  1 907.  Les  mollusques  fossiles  du  Tertiaire  et  du 
Cretace  Superieur  de  /'Argentine.  Anales  del  Museo  Na- 
cional de  Buenos  Aires,  ser.  3,  v.  7,  61 1 pp.,  18  pis. 

Illingworth,  J.F.  1 902.  The  anatomy  of  Lucapina  crenulata 
Gray.  Zoo/ogische  Jahrbucher,  Abtheilung  fur  Anatomie 
und  Ontogenie  der  Tiere  16:449-480,  pis.  31-33. 

Jara,  H.F.,  and  C. A.  Moreno.  1984.  Herbivory  and  struc- 
ture in  a midlittoral  rocky  community:  A case  in  south- 
ern Chile.  Ecology  65(  1):28— 38. 

Johnson,  R.I.  1964.  The  Recent  Mollusca  of  Augustus  Ad- 
dison Gould.  United  States  National  Museum,  Bulletin 
239,  182  pp.,  45  pis. 

Keen,  A.M.  1971.  Sea  shells  of  tropica!  west  America.  Sec- 
ond edition.  Stanford:  Stanford  University  Press,  xiv  + 
1064  pp.,  22  pis. 

Knight,  J.B.,  L.R.  Cox,  A.M.  Keen,  R.L.  Batten,  E.L.  Yoch- 
elson,  and  R.  Robertson.  1960.  Systematic  descrip- 
tions (Archaeogastropoda).  In  Treatise  on  invertebrate 
paleontology.  Part  I,  Mollusca  1.  ed.  R.C.  Moore,  1 69— 
310.  Geological  Society  of  America  and  University  of 
Kansas  Press. 


68  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


Lamarck,  J.B.P. A.  de  M.  de.  1815-1822.  Histoire  naturelle 
des  animaux  sans  vertebres.  Paris,  v.  1-7. 

. 1835-1845.  Histoire  naturelle  des  animaux  sans 

vertebres,  ed.  2,  revised  by  G.P.  Deshayes  and  H.  Milne- 
Edwards.  Paris,  v.  1-11. 

Lesson,  R.P.  1 830-183 1 . Voyage  autour  du  monde  . . . sur 
"la  Coquil/e” pendant . . . 1822-25.  Paris,  v.  2,  Zoologie, 
47 1 pp.,  atlas,  1 57  pis. 

Lindberg,  D.R.  1976.  The  homing  depression  of  the  limpet, 
Collisella  scabra  (Gould,  1846).  Bulletin  of  the  Ameri- 
can Malacological  Union  for  1975:69-70. 

Lindberg,  D.R.,  and  K.R.  Dwyer.  1983.  The  topography, 
formation  and  role  of  the  home  depression  of  Collisella 
scabra  (Gould)  (Gastropoda:  Acmaeidae).  The  Veliger 
25(3):229-234. 

Lowenstam,  H.A.  1954.  Factors  affecting  the  aragonite: 
calcite  ratios  in  carbonate-secreting  marine  organisms. 
Journal  of  Geology  62:284-322. 

. 1 964.  Coexisting  calcites  and  aragonites  from  skel- 
etal carbonates  of  marine  organisms  and  their  strontium 
and  magnesium  contents.  In  Recent  researches  in  the 
fields  of  hydrosphere,  atmosphere  and  nuclear  geochem- 
istry, editorial  committee  of  Saguawara  festival  volume, 
374-404.  Tokyo. 

MacClintock,  C.  1963.  Reclassification  of  gastropod  Pro- 
scutum Fischer  based  on  muscle  scars  and  shell  structure. 
Journal  of  Paleontology  37:141-156. 

. 1967.  Shell  structure  of  patelloid  and  bellerophon- 

tid  gastropods  (Mollusca).  Peabody  Museum  of  Natural 
History,  Yale  University,  Bulletin  22,  140  pp.,  32  pis. 

Margolin,  A. S.  1964.  The  mantle  response  of  Diodora  as- 
pera.  Animal  Behaviour  12(1):  187-1 94. 

Marincovich,  L.,  Jr.  1973.  Intertidal  mollusks  of  Iquique, 
Chile.  Natural  History  Museum,  Los  Angeles  County, 
Science  Bulletin  16,  49  pp. 

McLean,  J.H.  1984a.  A case  for  derivation  of  the  Fissu- 
rellidae  from  the  Bellerophontacea.  Malacologia  25(1): 
3-20. 

. 1984b.  Shell  reduction  and  loss  in  fissurellids:  A 

review  of  genera  and  species  in  the  Fissurellidea  group. 
American  Malacological  Bulletin  2:21-34. 

Melvill,  J.C.,  and  R.  Standen.  1898.  Notes  on  a collection 
of  marine  shells  from  Lively  Island,  Falklands,  with  list 
of  species.  Journal  of  Conchology  9:97-105. 

. 1907.  The  marine  Mollusca  of  the  Scottish  Na- 
tional Antarctic  Expedition.  Transactions  of  the  Royal 
Society  of  Edinburgh  46: 119-157. 

. 1914.  Notes  on  Mollusca  collected  in  the  north- 
west Falklands,  by  Mr.  Rupert  Vallentin,  F.L.S.,  with 
descriptions  of  six  new  species.  Annals  and  Magazine 
of  Natural  History,  series  8,  13:109-136. 

Mermod,  G.  1950.  Les  types  de  la  collection  Lamarck  au 
Museum  de  Geneve.  Mollusques  vivantes,  I.  Revue  Suisse 
de  Zoologie  57(34):687-756. 

Metivier,  B.  1969.  Resultats  scientiftques  des  compagnes 
de  la  “Calypso.”  Campagne  au  large  des  cotes  Atlan- 
tiques  de  I’Amerique  du  Sud  (1961-1962).  Mollusques 
Prosobranches:  Fissurellidae,  Acmaeidae  et  Patellidae. 


Annales  de  TInstitut  Oceanographique,  Monaco  47(1): 

1 15-125. 

Moreno,  C.A.,  and  E.  Jaramillo.  1983.  The  role  of  grazers 
in  the  zonation  of  intertidal  macroalgae  of  the  Chilean 
coast  near  Valdivia.  Oikos  41:73-76. 

Moreno,  C.A.,  J.P.  Sutherland,  and  H.F.  Jara.  1984.  Man 
as  a predator  in  the  intertidal  zone  of  southern  Chile. 
Oikos  42:155-160. 

Odhner,  N.H.  1932.  Zur  Morphologie  und  Systematik  der 
Fissurelliden.  Jenaische  Zeitschrift  fur  Naturwissen- 
schaft  67:292-309. 

Olsen,  D.A.  1968a.  Banding  patterns  of  Haliotis  rufescens 
as  indicators  of  botanical  and  animal  succession.  Bio- 
logical Bulletin  37(  1 ):  1 39—14 1 . 

. 1968b.  Banding  patterns  in  Haliotis  II.  Some  be- 
havioral considerations  and  the  effect  of  diet  on  shell 
coloration  for  H.  rufescens,  H.  corrugata,  H.  sorenseni, 
and  H.  assimilis.  The  Veliger  1 1(2):  1 35-1 39. 

Orbigny,  A.d’.  1831-1847.  Voyage  dans  I’Amerique  Mer- 
idional. Paris,  v.  5,  part  3,  pp.  1-758;  atlas,  v.  9,  Moll- 
usques, pis.  1-85. 

Paine,  R.T.,  and  A. R.  Palmer.  1978.  Sicyases  sanguineus: 
A unique  trophic  generalist  from  the  Chilean  intertidal 
zone.  Copeia  1978(1):7 5—8 1 . 

Pena,  G.M.  1970.  Zonas  de  distribucion  de  los  gastero- 
podos  marinos  del  Peru.  Anales  Cientificos  de  la  Uni- 
versidad  Nacional  Agraria  (Lima)  8(3-4):  1 53-170. 

Perez-Farfante,  I.  1943.  The  genera  Fissurella,  Lucapina, 
and  Lucapinella  in  the  western  Atlantic.  Johnsonia  1(10): 
1-20. 

. 1952.  Nuevo  genero  y nuevo  especie  del  genero 

Fissurella.  Revista  de  la  Sociedad  Malacologica  "Carlos 
de  la  Torre”  (Habana)  9(1): 3 1-3 3. 

Philippi,  R. A.  1845.  Diagnosen  einiger  neuer  Conchylien. 
Archiv  fur  Naturgeschichte  1 1(1): 50—7 1 , 142-144. 

. 1845-1846.  Fissurella.  Abbildungen  und  Beschrei- 

bungen  neuer  oder  wenig  bekannter  Conchylien.  Kassel, 
2 pis.  [pi.  1,  1845;  pi.  2,  1846], 

. 1857.  Ueber  die  Conchylien  der  Magellanstrasse. 

Malakozoologische  Blatter  3: 1 57-173. 

. 1860.  Reise  durch  die  Wueste  Atacama  auf  Befehl 

der  Chilenischen  Regierung  im  Sommer  1853-54.  Halle: 
E.  Anton,  254  pp.,  27  pis.  [Also  in  Spanish  edition  ( 1 860) 
“ Viage  al  Desierto  de  Atacama  . . . .”] 

. 1887.  Die  Tertiaren  und  Quartaren  Versteinerun- 

gen  Chiles.  Leipzig,  268  pp.,  58  pis.  [Also  in  Spanish 
edition  (1887),  Santiago.] 

Pilsbry,  H.A.  1890.  Stomatellidae,  Scissurellidae,  Pleuro- 
tomariidae,  Haliotidae,  Scutellinidae,  Addisoniidae, 
Cocculinidae,  Fissurellidae.  Manual  of  Conchology,  v. 
12.  Philadelphia. 

Powell,  A.  W.B.  1951.  Antarctic  and  subantarctic  Mollusca: 
Pelecypoda  and  Gastropoda.  Discovery  Reports  26:47- 
196,  pis.  5-10. 

Ramirez-Boehme,  J.  1974.  Nuevas  especies  chilenas  de 
Lucapina,  Fissurella,  y Collisella  (Mollusca,  Archaeo- 
gastropoda).  Boletin  Museo  Nacional  de  Historia  Nat- 
ural, Chile  33:15-34. 


Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella  69 


Ringuelet,  R.A.,  A.  Amor,  N.  Magildi,  and  R.  Pallares. 
1962.  Estudio  ecologico  de  la  fauna  intercotidal  de 
Puerto  Deseado,  en  Febrero  de  1 96 1 (Santa  Cruz,  Argen- 
tina). Physis  (Buenos  Aires)  23(64):35-53. 

Rathke,  M.H.  1833.  Zoologischer  Atlas.  Part  5.  Berlin,  i- 
vii,  1-28,  pis.  21-25.  [Completion  of  work  started  by 
J.F.  Eschscholtz,  1829-1831.] 

Reeve,  L.  1849-1850.  Monograph  of  the  genus  Fissurella. 
Conchologica  lconica.  London,  v.  6,  16  pis.  [pis.  1-8, 
1849;  pis.  9-16,  1850], 

Riveros-Zuniga,  F.  1951.  Catalogo  descriptivo  de  fisure- 
lidos  Chilenos.  Revista  de  Biologfa  Marina  (Universidad 
de  Chile)  3(1— 2):89— 148. 

Rochebrune,  A.T.  de,  and  J.  Mabille.  1885.  Diagnoses  de 
Mollusques  nouveaux,  receuillis  par  les  membres  de  la 
mission  du  Cap  Horn  ...  a Santa  Cruz.  Bulletin  de  la 
Societe  Phi/omathique  de  Paris  7(9):  1 00-1 1 1 . 

. 1889.  Mollusques.  Mission  Scientifique  du  Cap  Horn 

(1882-1883),  v.  6,  Zoologie.  Paris,  126  pp.,  8 pis. 

Scarabino,  V.  1977.  Moluscos  del  Golfo  San  Matias  (Pro- 
vincia  de  Rio  Negro,  Republica  Argentina).  Comuni- 
caciones  de  !a  Sociedad  Malacologica  del  Uruguay  4(3 1 — 
32):  1 77—297. 

Sowerby,  G.B.  1825.  A catalogue  of  the  shells  contained  in 
the  collection  of  the  late  Earl  ofTankerville.  London,  vii 
+ 99  + xxxiv  pp.,  9 pis. 

. 1835a.  Shells  collected  by  Mr.  Cuming  on  the  west- 
ern coast  of  South  America,  and  among  the  islands  of 
the  South  Pacific  Ocean.  Proceedings  of  the  Zoological 
Society  of  London,  for  1 834: 1 23-1 28. 

. 1835b.  A catalogue  of  the  Recent  species  of  Fissu- 
rella. The  conchologica!  illustrations.  London,  pp.  1-8, 
77  figs,  on  12  unnumbered  plates. 

Sowerby,  G.B.,  II.  1 862.  Monograph  of  the  family  Fissu- 
rellidae,  genus  Fissurella.  Thesaurus  conchyliorum,  or 
Monographs  of  genera  of  shells.  London,  v.  3,  pp.  183- 
206,  pis.  1-9. 

Steams,  R.E.C.  1891.  List  of  shells  collected  on  the  west 
coast  of  South  America,  principally  between  latitudes 
7°30'  S.  and  8°49'  N.,  by  Dr.  W.H.  Jones,  surgeon,  U.S. 
Navy.  Proceedings  of  the  United  States  National  Mu- 
seum 14(854):307-335. 

Strebel,  H.  1907.  Beitrage  zur  Kenntnisder  Mollusken  Fau- 
na der  Magalhaen-Provinz.  5.  Zoologische  Jahrbiicher 


Abteilung  fur  Systematik,  Geographic  und  Biologic,  Jena 
25:79-196. 

. 1908.  Die  Gastropoden.  Wissenschaftliche  Ergeb- 

nisse  der  Schwedische  Sudpolar-Expedition,  1901-1903, 
v.  6,  Zoologie  2.  Stockholm,  1 1 1 pp. 

Stuardo,  J.  1964.  Distribucion  de  los  moluscos  marinos 
litorales  en  latinoamerica.  Boletin  Instituto  Biologia 
Marina  (Mar  del  Plata ) 7:79-91. 

Tobler,  M.  1902.  Zur  Anatomie  von  Parmophorus  inter- 
medius  Reeve.  Jenaische  Zeitschrift  fur  Naturwissen- 
schaft  36:229-274,  pis.  13-15. 

Tomlinson,  J.T.  1969.  The  burrowing  barnacles  (Cirripe- 
dia:  Order  Acrothoracica).  United  States  National  Mu- 
seum, Bulletin  296,  162  pp. 

Vance,  R.S.  1978.  A mutualistic  interaction  between  a ses- 
sile marine  clam  and  its  epibionts.  Ecology’  59(4):679- 
685. 

Vermeij,  G.J.  1978.  Biogeography  and  adaptation.  Cam- 
bridge: Harvard  University  Press,  xi  + 332  pp. 

Ward,  J.  1966a.  Feeding,  digestion,  and  histology  of  the 
digestive  tract  in  the  keyhole  limpet  Fissurella  barba- 
densis  Gmelin.  Bulletin  of  Marine  Science  16:668-684. 

. 1966b.  The  breeding  cycle  of  the  keyhole  limpet 

Fissurella  barbadensis  Gmelin.  Bulletin  of  Marine  Sci- 
ence 16:685-695. 

Watson,  R.B.  1886.  Report  on  Scaphopoda  and  Gastero- 
poda  collected  by  “Challenger.  ” Report  on  the  Scientific 
Results  of  the  Voyage  of  the  H.M.S.  Challenger.  Zool- 
ogy, v.  15.  London,  vii  + 756  pp.,  50  pis. 

Wenz,  W.  1938-1944.  Gastropoda.  Allgemeiner  Teil  und 
Prosobranchia.  In  Handbuch  der  Palaozoologie,  ed.  O.H. 
Schindewolf.  Band  6.  Berlin,  1639  pp. 

Woodring,  W.P.  1965.  Endemism  in  Middle  Miocene  Ca- 
ribbean molluscan  faunas.  Science  148:961-963. 

. 1966.  The  Panama  land  bridge  as  a sea  barrier. 

Proceedings  of  the  American  Philosophical  Society 
1 1 0(6):425— 433. 

Ziegenhom,  A.,  and  H.  Thiem.  1925.  Beitrage  zur  Syste- 
matik und  Anatomie  der  Fissurellen.  Jenaische  Zeit- 
schrift fur  Naturwissenschaft  62:1-78,  pis.  1-6. 


Submitted  25  October  1983;  accepted  for  publication  3 May 
1984. 


70  Contributions  in  Science,  Number  354 


McLean:  Peruvian  and  Magellanic  Fissurella 


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Q 

11 

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Number  355 
6 December  1984 


AN  EARLY  HEMINGFORDIAN  (EARLY  MIOCENE) 
FOSSIL  VERTEBRATE  FAUNA  FROM  BORON, 


WESTERN  MOJAVE  DESERT,  CALIFORNIA 


David  P.  Whistler 


Natural  History  Museum  of  Los  Angeles  County  • 900  Exposition  Boulevard  • Los  Angeles,  California  90007 


.'•i 


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Printed  at  Allen  Press,  Inc.,  Lawrence,  Kansas 


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AN  EARLY  HEMINGFORDIAN  (EARLY  MIOCENE) 
FOSSIL  VERTEBRATE  FAUNA  FROM  BORON, 
WESTERN  MOJAVE  DESERT,  CALIFORNIA 

David  P.  Whistler1 


ABSTRACT.  A small  assemblage  of  fossil  vertebrates  has  been  ob- 
tained from  the  Arkose  Member  of  the  Kramer  Beds  in  the  upper 
part  of  the  Tropico  Group  at  Boron,  California.  Known  as  the  Boron 
Local  Fauna  (new  name),  it  displays  affinities  with  the  Tick  Canyon 
and  Phillips  Ranch  Local  Faunas  of  southern  California  and  the 
fossil  assemblages  from  the  Runningwater  and  Batesland  Formations 
of  Nebraska  and  South  Dakota,  respectively,  the  latter  used  as  the 
standard  for  the  earlier  part  of  the  Hemingfordian  land  mammal 
age.  These  correlations  and  radiometric  dates  with  a range  of  18.3 
± 0.6  ma  to  20.3  ± 0.7  ma  from  the  underlying  Saddleback  Basalt 
support  an  early  Hemingfordian  age  for  the  Boron  Local  Fauna. 

Prior  to  the  discovery  of  this  fossil  assemblage,  considerable  un- 
certainty existed  about  the  age  of  scattered  occurrences  of  volcanic 
flows  and  volcaniclastic  rocks  in  the  western  Mojave  Desert  which 
were  restricted  and  redefined  as  the  Tropico  Group  by  Dibblee 
(1958a).  Based  on  lithologic  correlations,  ages  ranging  from  Middle 
Miocene  to  Early  Pliocene  had  been  suggested  for  the  T ropico  Group. 
The  presence  of  the  early  Hemingfordian  Boron  Local  Fauna  in  the 
uppermost  unit  of  the  Tropico  Group  limits  the  age  of  this  entire 
sequence  to  pre-Middle  Miocene.  The  well-preserved  assemblage 
contains  at  least  15  vertebrate  taxa.  Two  new  species  are  described, 
Cupidinimus  boronensis  n.  sp.  (Rodentia,  Heteromyidae)  and  Ale- 
tomeryx  occidentalis  n.  sp.  (Artiodactyla,  Palaeomerycidae).  There 
are  also  two  lizards  which  are  not  diagnostic  at  the  family  level,  two 
snakes,  cf.  Calamagras  sp.  (Boidae)  and  cf.  Paracoluber  sp.  (Colu- 
bridae),  a lagomorph,  Archaeolagus  sp.  or  Hypolagus  sp.  (Leporidae), 
three  additional  rodents,  IMiospermophilus  sp.  (Sciuridae),  Moo- 
komys  sp.  and  Trogomys  cf.  T.  rupinimenthae  (Heteromyidae)  and 
five  additional  artiodactyls,  Merychyus  cf.  M.  minimus  and  IMer- 
ychyus  sp.  (Merycoidodontidae),  cf.  Hesperocamelus  sp.  (Camelidae) 
and  two  other  camels  not  identified  to  genus.  Conspicuous  by  their 
absence  are  eomyid  and  large  geomyoid  rodents,  horses,  dromo- 
merycine  palaeomerycids,  and  antilocaprids  typical  of  most  Middle 
Miocene  assemblages. 

INTRODUCTION 

In  March,  1964,  the  geology  staff  of  the  United  States  Borax 
and  Chemical  Corporation  mine  at  Boron,  California,  re- 
ported uncovering  a pocket  of  fossil  bones  during  operations 


in  the  open  pit  mine  (Figure  1).  An  initial  sample  collected 
by  the  Department  of  Geology  at  the  University  of  California 
at  Riverside  was  recognized  as  representing  a Miocene  as- 
semblage older  than  any  previously  known  from  the  central 
Mojave  Desert.  The  fossils  were  recovered  from  the  Kramer 
Beds,  sediments  which  were  correlated  with  other  scattered 
continental  sequences  in  the  Mojave  Desert,  all  of  which  were 
considered  to  be  Late  Miocene  or  Pliocene  in  age.  These 
fossil  vertebrates,  hereafter  called  the  Boron  Local  Fauna, 
demonstrated  that  the  Kramer  Beds  were  older  than  the  rich- 
ly fossiliferous  deposits  of  the  Barstow  Formation  which  con- 
tained the  oldest  fossil  faunas  then  known  from  the  central 
Mojave  Desert  and  necessitated  re-thinking  the  geologic  his- 
tory of  this  area.  Subsequent  discoveries  (Woodbume  et  al., 
1974;  Woodbume,  Miller,  and  Tedford,  1982)  have  led  to 
recognition  of  even  earlier  Miocene  assemblages,  but  the 
Boron  Local  Fauna  was  the  first  well-preserved  pre-Barsto- 
vian  assemblage  from  the  Mojave  Desert. 

The  Boron  Local  Fauna  is  markedly  different  in  taxonomic 
composition  from  the  well-documented  faunas  of  the  Bar- 
stow Formation  and  other  fairly  widespread  Barstovian 
(Middle  Miocene)  assemblages  in  southern  California.  The 
Boron  Local  Fauna  is  dominated  by  four  groups  (heteromyid 
rodents,  oreodonts,  camels,  and  palaeomerycids)  which,  ex- 
cept for  camels,  are  rare  or  absent  in  Barstovian  assemblages 
of  southern  California.  In  contrast,  the  Boron  Local  Fauna 
lacks  three  groups  (cricetid  rodents,  horses,  and  antiloca- 
prids) which  are  a common  element  of  Barstovian  assem- 
blages. The  Boron  Local  Fauna  also  differs  from  the  less  well 
preserved  late  Hemingfordian  assemblages  of  southern  Cal- 
ifornia which  are  dominated  by  small  merychippine  horses, 
middle-sized  oreodonts,  dromomerycines,  and  antilocaprids, 
groups  lacking  at  Boron.  The  Boron  Local  Fauna  appears  to 


1 Vertebrate  Paleontology  Section,  Natural  History  Museum  of 
Los  Angeles  County,  900  Exposition  Blvd.,  Los  Angeles,  California, 
90007. 


Contributions  in  Science,  Number  355,  pp.  1-36 
Natural  History  Museum  of  Los  Angeles  County,  1984 


ISSN  0459-8113 


118 


117 


Figure  1.  Index  map  showing  location  of  Boron  Local  Fauna  and  other  southern  California  Miocene  fossil  vertebrate  assemblages  discussed 
in  this  report. 


be  the  first  early  Hemingfordian  fossil  assemblage  sampled 
in  southern  California. 

GEOLOGIC  SETTING 

The  western  and  central  Mojave  Desert  is  a broad,  fault- 
bounded  block  underlain  by  Mesozoic  crystalline  rocks  which 
were  faulted  into  many  localized  basins  during  the  Middle 
and  Late  Cenozoic.  Deposition  in  these  basins  was  domi- 


nated by  volcanics  and  volcaniclastics  during  the  first  half 
of  the  Miocene.  Volcanism  subsided  during  later  Miocene 
and  fluviatile  and  lacustrine  sediments  accumulated  in  the 
localized  basins  which  persisted  due  to  down-faulting,  a pro- 
cess which  continues  today  in  some  areas.  In  other  areas, 
faulting  has  uplifted  and  exposed  Miocene  and  younger  se- 
quences. 

Considerable  geologic,  stratigraphic,  and  geophysical  work 
has  been  done  in  the  general  area  of  Boron,  much  of  it  per- 


2 Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna 


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Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna  3 


Figure  2.  Stratigraphic  nomenclature  of  geologic  units  at  Boron  and  vicinity. 


taining  to  the  deposition  and  extent  of  the  borate-rich  sed- 
iments there  (Figure  2).  The  Tertiary  rocks  at  Boron  and 
surrounding  hills  were  originally  referred  to  as  the  Rosamond 
Series  (Hershey,  1902),  a name  expanded  to  include  most  of 
the  continental  sedimentary  and  volcanic  rocks  in  the  Mo- 
jave Desert,  including  such  units  as  the  Ricardo  Formation 
of  Clarendonian  age  (Merriam,  1919;  Dibblee,  1952)  and 
widely  scattered  deposits  of  Hemingfordian  and  Barstovian 
age,  usually  referred  to  the  Barstow  Formation  (Merriam, 
1919;  Bowen,  1954).  The  first  mapping  of  the  borate  pro- 
ducing beds  at  Boron  was  done  by  Gale  (1946).  He  referred 
these  beds  to  the  Ricardo  Formation  and  defined  a new  un- 
derlying unit,  the  Saddleback  Basalt,  which  occurs  in  surface 
exposures  at  Saddleback  Butte,  5 km  to  the  east  of  Boron. 
A similar  basaltic  flow  also  occurs  as  a series  of  small  hills 
just  north  of  the  mine  and  is  found  exposed  in  several  areas 
of  the  open  pit.  It  is  also  recognized  in  many  of  the  drill  holes 
in  the  general  area. 

During  the  1950’s,  Dibblee  undertook  a regional  mapping 
program  in  the  western  and  central  Mojave  Desert  including 
the  area  around  Boron.  Because  of  confusion  over  previous 
usage  of  stratigraphic  names,  Dibblee  (1958a)  established  a 
new  unit,  the  Tropico  Group,  to  include  only  some  of  the 
Tertiary  rocks  previously  referred  to  as  the  Rosamond  Series. 
He  included  four  formations  within  the  Tropico  Group  (Gem 
Hill  Formation,  Fiss  Fanglomerate,  Bissel  Formation,  and 
Saddleback  Basalt),  but  excluded  such  units  as  the  Ricardo 
and  Barstow  Formations  which  had  been  included  in  the 
Rosamond  Series  by  other  workers. 

Dibblee  (1958a,  b)  did  not  give  any  formational  designa- 
tion to  the  beds  at  the  open  pit  mine  at  Boron  but  referred 
to  them  only  as  the  “Upper  Part  of  the  Tropico  Group.” 
Following  the  usage  of  the  geology  staff  at  the  Boron  mine, 
Bowser  (1965),  in  his  geochemical  studies,  used  the  term 
“Kramer  Lake  Beds”  for  the  sequence  above  the  Saddleback 
Basalt.  Barnard  and  Kistler  (1966)  simplified  Bowser’s  name 
to  the  Kramer  Beds,  included  the  Saddleback  Basalt  and 
published  a detailed  stratigraphic  section  which  included  the 
location  of  the  Boron  Local  Fauna.  They  subdivided  the 
Kramer  Beds  into  three  members,  the  Saddleback  Basalt, 
Shale  Member,  and  Arkose  Member  in  ascending  order.  The 
Saddleback  Basalt  forms  the  base  of  the  exposed  section  in 
the  open  pit  mine  and  is  the  only  member  of  the  Kramer 
Beds  with  natural  exposures.  This  sequence  of  flows  and  flow 
breccias  appears  to  lie  unconformably  on  pyroclastic  rocks 
which  may,  in  part,  represent  an  eastern  extension  of  the 
Gem  Hill  Formation,  the  basal  unit  of  the  Tropico  Group. 

The  Shale  and  Arkose  Members  of  the  Kramer  Beds  over- 
lying  the  Saddleback  Basalt  include  lacustrine  and  fluviatile 
sediments  exposed  only  in  the  open  pit  mine  and  known 
otherwise  from  drilling  data  (Benda  et  al.,  1960;  Dickey, 
1957).  The  seemingly  local  extent  of  these  units  has  been 
discussed  by  several  authors  (Gale,  1946;  Bowser,  1964;  Bar- 
nard and  Kistler,  1966).  The  Shale  Member  is  lacustrine, 
composed  of  dark  green  to  brown,  borate-bearing  clays,  shales, 
stratified  borate  ore  bodies,  and  several  thin  bentonitic  tuff 
beds.  The  Arkose  Member,  which  produced  the  Boron  Local 


Fauna,  is  composed  of  beds  of  well-bedded,  buff,  light  tan 
and  dark  reddish  brown  claystone  and  micaceous  siltstone 
and  lenses  of  coarser,  cross-bedded,  light  brown  to  yellowish 
white  arkosic  sandstone  and  pebble  conglomerate.  The  Ar- 
kose Member  contains  relatively  few  volcanic  beds,  sug- 
gesting that  the  volcanism  so  commonly  represented  in  the 
underlying  units  of  the  Tropico  Group  had  subsided. 

At  the  time  of  his  original  description,  Dibblee  (1958a) 
believed  that  much  of  the  Tropico  Group  was  Late  Miocene 
and  Pliocene  in  age.  His  conclusion  was  based  on  a diatom 
flora  recovered  from  a limestone  in  the  Tropico  Group  west 
of  Boron  and  on  Gale’s  ( 1 946)  lithologic  correlation  of  the 
Saddleback  Basalt  with  andesite  breccia  flows  in  the  basal 
part  of  the  Ricardo  Formation  which  produced  a fossil  ver- 
tebrate fauna  of  Clarendonian  age  (then  considered  Pliocene). 
The  discovery  of  the  Boron  Local  Fauna  at  first  seemed  to 
refute  this  latter  conclusion,  but  subsequent  work  in  the  type 
area  of  the  Ricardo  Formation  has  shown  that  a correlation 
of  the  basal  andesite  breccia  flows  and  underlying  pyroclastic 
rocks  with  the  Tropico  Group  is  reasonable.  Field  studies 
have  shown  that  these  volcanic  rocks  unconformably  un- 
derlie the  fossil-producing  rocks  of  the  Ricardo  Formation 
which  are  radiometrically  dated  at  a maximum  of  10.2  ma 
(Evemden  et  al.,  1964).  These  volcanics  have  subsequently 
been  radiometrically  dated  at  17.1  ma  (Loomis  et  al.,  1983) 
and  thus  fall  within  the  age  range  of  the  Tropico  Group.  The 
Saddleback  Basalt  has  yielded  radiometric  dates  in  a range 
from  18.3  ± 0.6  (R.B.  Kistler,  personal  communication)  to 
20.3  ± 0.7  ma  (Armstrong  and  Higgens,  1973),  somewhat 
older  than,  but  possibly  representing  the  same  volcanic  ep- 
isode as  the  andesite  flows  underlying  the  Ricardo  Forma- 
tion. 

Dibblee  (1958a)  recognized  lithologic  similarities  between 
the  Gem  Hill  Formation,  the  basal  unit  of  the  Tropico  Group, 
the  Middle  Miocene  Kinnick  Formation  in  the  mountains 
north  of  Tehachapi  and  the  Pick  Handle  Formation  under- 
lying the  Middle  and  Late  Miocene  Barstow  Formation  in 
the  Mud  and  Calico  Hills.  The  presence  of  the  Boron  Local 
Fauna  in  the  uppermost  member  of  the  Tropico  Group  tends 
to  support  Dibblee’s  conclusions,  but  also  shows  that  this 
entire  unit  is  pre-Middle  Miocene  in  age,  much  older  than 
he  had  supposed. 

FOSSIL  OCCURRENCE 

Fossils  of  the  Boron  Local  Fauna  were  not  observed  at  the 
original  site  of  deposition  by  the  author.  They  were  first 
noticed  by  several  mine  employees  operating  heavy  equip- 
ment removing  overburden  from  the  open  pit  mine.  The 
mine  foreman  informed  the  geology  staff  who  had  the  re- 
maining fossil-bearing  matrix  dumped  in  a separate  location 
from  the  general  mine  dump  so  that  the  fossils  could  be 
recovered.  It  is  not  certain  how  much  of  the  fossil-bearing 
pocket  was  lost  before  this  action  was  taken,  but  4-5  earth- 
mover  loads  (12-15  cubic  meters)  were  subsequently  re- 
covered before  the  pocket  stopped  producing  readily  visible 
fossil  bone. 


4 Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna 


According  to  the  location  provided  by  mine  employees, 
the  fossil-producing  layer  was  approximately  146  m above 
the  Saddleback  Basalt.  Observations  made  in  the  pit  in  1 964 
and  from  the  fossil-bearing  matrix  indicate  the  fossils  were 
restricted  to  one  or  more  beds  of  light  tan  micaceous  clay- 
stones  which  locally  contain  pumice  lapilli  fragments.  These 
beds  were  part  of  a sequence  of  fluviatile  siltstones  and  sand- 
stones locally  cut  by  channels  of  coarser  arkosic  sandstones 
and  pebble  conglomerates.  About  3 m above  the  fossil-pro- 
ducing beds  were  several  discontinuous  lenses  of  grayish- 
pink  bentonitic  tuff  and  yellowish-white,  medium-grained 
sandy  tuff. 

Many  of  the  individual  bones  were  fractured  and  splintered 
prior  to  fossilization,  suggesting  a period  of  surface  exposure 
before  burial.  On  the  other  hand,  some  fossils  are  very  well 
preserved  and  several  limb  fragments  were  still  partially  ar- 
ticulated. There  is  no  evidence  of  rodent  or  carnivore  gnaw- 
ing on  any  of  the  fossil  bone.  Several  matrix  blocks  contain 
small,  lenticular  concentrations  of  partially  leached  small 
vertebrate  bone,  apparently  representing  disassociated  owl 
pellets  or  carnivore  coprolites  which  undoubtedly  are  the 
source  of  the  rich  microvertebrate  assemblage. 

Limited  prospecting  by  the  author  in  the  walls  of  the  open 
pit  mine  in  1964  did  not  produce  additional  fossils,  nor  did 
extensive  prospecting  in  all  exposed  fluviatile  facies  of  the 
Tropico  Group  within  15  km  of  Boron  yield  further  fossil 
specimens.  The  entire  collection  of  fossil  vertebrates  was 
deposited  in  the  Department  of  Earth  Sciences  at  the  Uni- 
versity of  California  at  Riverside. 

METHODS 

The  fossils  were  obtained  from  the  pile  of  matrix  removed 
from  the  open  pit  mine.  Blocks  containing  natural  molds  of 
bones  destroyed  during  excavation  were  also  collected  and 
plaster  casts  were  subsequently  made  of  these  molds.  All 
matrix  with  a lithology  similar  to  known  fossil  bearing  matrix 
(about  2700  lb)  was  sieved  under  water  with  screens  with  an 
average  mesh  opening  of  0.7  mm. 

Measurements  on  larger  bones  were  made  with  a vernier 
caliper  to  the  nearest  0. 1 mm  and  the  microvertebrates  were 
measured  with  an  EPOI  optical  micrometer  to  the  nearest 
0.01  mm.  Measurements  are  taken  at  their  maximum  di- 
mensions unless  otherwise  indicated.  The  anteroposterior 
dimension  of  the  P4  of  heteromyids  was  measured  with  the 
posterior  surface  of  the  metaloph  oriented  vertically. 

The  stereophotographs  were  taken  by  the  author  using  an 
extension  bellows  and  reversed  50  mm,  f 1.4  lens  mounted 
on  a stereophotographic  bar.  The  specimens  were  coated  with 
vaporized  magnesium  oxide  prior  to  photographing. 

Radiometric  dates  have  been  converted  using  the  new  de- 
cay and  abundance  constants  given  by  Dalrhymple  (1979). 
This  has  the  effect  of  increasing  most  later  Tertiary  dates 
published  before  1979  by  about  2.6  percent. 

Metric  abbreviations  and  designations  of  tooth  position 
follow  standard  usage.  Other  abbreviations  used  are  as  fol- 
lows: 


LACM 

Collections  of  the  Natural  History  Museum  of 
Los  Angeles  County 

LACM(CIT)  Collections  formerly  at  the  California  Institute 
of  Technology,  now  at  LACM 

UCMP 

Collections  of  the  University  of  California, 
Museum  of  Paleontology,  Berkeley 

UCR 

Collections  of  the  University  of  California  at 
Riverside,  Department  of  Geological  Sciences 

L 

Left 

R 

Right 

ma 

millions  of  years  before  present 

FAUNAL  LIST 

Below  is  a summary  of  the  fossil  vertebrate  assemblage  of 
the  Boron  Local  Fauna  as  recognized  in  the  following  sys- 
tematics  section. 

Class  Reptilia 
Order  Squamata 
Suborder  Sauria 

Family  Iguanidae  or  Xantusiidae 

Iguanidae  or  Xantusiidae  sp.  indet. 

Family  undetermined 
Sauria,  incertae  sedis 
Suborder  Serpentes 
Family  Boidae 

Subfamily  Erycinae 
cf.  Calamagras  sp. 

Family  Colubridae 
cf.  Paracoluber  sp. 

Class  Mammalia 
Order  Lagomorpha 
Family  Leporidae 
Subfamily  Archaeolaginae 
Arachaeolagus  sp.  or  Hypolagus  sp. 

Order  Rodentia 

Family  Sciuridae 

IMiospermophilus  sp. 

Family  Heteromyidae 
Subfamily  Perognathinae 
Mookomys  sp. 

Trogomys  cf.  T.  rupinimenthae 
Subfamily  Dipodomyinae 
Cupidinimus  boronensis  n.  sp. 

Order  Artiodactyla 

Family  Merycoidodontidae 
Subfamily  Merychinae 

Merychyus  cf.  M.  minimus 
IMerychyus  sp. 

Family  Camelidae 

Subfamily  Aepycamelinae 
cf.  Hesperocamelus  sp. 

Subfamily  undetermined 
Camelidae,  small  species 
Camelidae,  large  species 


Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna  5 


Family  Palaeomerycidae 
Subfamily  Aletomerycinae 
Aletomeryx  occidentalis  n.  sp. 

SYSTEMATICS 
Class  Reptilia 
Order  Squamata 

Lizards  and  snakes  are  represented  by  very  fragmentary  ma- 
terial, but  the  presence  of  two  lizards  and  two  snakes  can  be 
demonstrated.  The  lizard  identifications  are  based  on  the 
tooth-bearing  bones  although  some  postcranial  material  is 
also  present.  One  of  the  lizards  may  represent  the  only  Middle 
Miocene  record  of  the  family. 

At  least  one  boid  and  one  colubrid  snake  are  present.  Small 
boids  are  common  in  the  middle  and  late  Tertiary,  thus  the 
Boron  occurrence  is  not  unusual.  On  the  other  hand,  the 
colubrid  represents  one  of  the  earliest  records  of  the  family 
in  North  America  and  extends  the  geographic  range  of  such 
colubrids  to  west  of  the  Rocky  Mountains.  Vertebral  ter- 
minology follows  Auffenberg  (1963)  and  taxonomy  follows 
Holman  (1979). 

Suborder  Sauria 

Family  Iguanidae  or  Xantusiidae 
Iguanidae  sp.  indet.  or  Xantusiidae  sp.  indet. 

MATERIAL.  Fragment  of  right  dentary  (or  splenio-den- 
tary),  UCR  21 174;  anterior  left  dentary  (or  splemo-dentary), 
UCR  21175;  premaxilla  fragment,  UCR  21176;  miscella- 
neous unnumbered  edentulous  maxillary  and  dentary  frag- 
ments. 

DESCRIPTION.  Small  lizard,  dentary  less  than  1 cm  long 
based  on  estimate  from  several  specimens;  Meckelian  groove 
closed  and  fused  except  for  anterior  opening;  posterior  teeth 
strongly  tricuspate  with  grooves  on  both  lingual  and  labial 
tooth  surfaces  between  main  cusp  and  lateral  cusps;  anterior 
teeth  single  cusped;  teeth  of  similar  width  along  entire  tooth 
row;  subdental  lingual  shelf  well  developed;  dentary  thin 
(dorsoventrally)  below  subdental  lingual  shelf;  dentary  same 
thickness  below  subdental  lingual  shelf  for  most  of  length  of 
tooth  row;  premaxilla  with  6 teeth. 

DISCUSSION.  The  Boron  lizard  material  is  very  frag- 
mentary and  the  above  description  is  based  on  a composite 
of  several  specimens  with  the  single  common  character  of  a 
fused  Meckelian  canal,  a character  restricted  to  the  xantu- 
siids,  gekkonids,  and  some  iguanids  such  as  Dipsosaurus  Hal- 
lowell,  1854.  The  material  is  too  incomplete  to  precisely 
estimate  the  entire  tooth  row  length  or  the  number  of  teeth. 
The  individual  tooth  bases  are  relatively  large  and  uncrowd- 
ed, typical  of  xantusiids  and  some  iguanids  and  in  contrast 
to  gekkonids,  which  usually  have  large  numbers  of  small 
teeth.  No  posterior  dentary  fragments  are  preserved,  thus  the 
presence  or  absence  of  the  diagnostic  xantusiid  character  of 
separate  anterior  inferior  alveolar  and  anterior  mylohyoid 
foramina  (terminology  of  Schatzinger,  1980)  entirely  en- 


closed within  the  splenio-dentary  is  not  preserved.  The  teeth 
are  strongly  tricuspate,  similar  to  Paleoxantusia  kyrentos 
Schatzinger,  1980,  but  less  so  than  in  living  Dipsosaurus  or 
Xantusia  riversiana  Cope,  1883.  The  subdental  lingual  shelf 
is  well  developed  as  in  the  extinct  Paleoxantusia  Hecht,  1956, 
and  in  contrast  to  most  iguanids.  The  continuously  dorso- 
ventrally thin  dentary  below  the  subdental  lingual  shelf  is 
more  similar  to  that  of  xantusiids  than  iguanids  or  gekkonids. 

The  Boron  material  does  not  permit  positive  identification, 
but  appears  to  represent  an  undescribed  species  of  either 
xantusiid  or  iguanid.  Small  iguanids  are  poorly  known  in  the 
fossil  record  but  a variety  of  undescribed  material  is  present 
in  Middle  and  Late  Miocene  deposits  of  the  Mojave  Desert 
and  Great  Basin  (collections  at  the  University  of  California, 
Berkeley,  Museum  of  Paleontology,  Natural  History  Mu- 
seum of  Los  Angeles  County  and  University  of  California  at 
Riverside).  Paleoxantusia,  an  early  xantusiid,  has  been  rec- 
ognized only  from  the  Paleocene  and  Eocene  of  western  North 
America.  Undescribed  xantusiid  material  is  known  from 
Middle  and  Late  Miocene  deposits  in  the  Mojave  Desert 
(specimens  in  same  collections  as  above)  and  the  Middle 
Pleistocene  Palm  Springs  Formation  in  San  Diego  and  Im- 
perial Counties  of  California  (M.  Norell,  personal  commu- 
nication; specimens  in  the  Natural  History  Museum  of  Los 
Angeles  County). 

Family  undetermined 
Sauria  incertae  sedis 

DISCUSSION.  There  is  a single  dentary  fragment,  UCR 
2 1 1 77,  and  miscellaneous  unnumbered  specimens  of  a small 
lizard  with  an  unfused  Meckelian  canal.  The  tooth  mor- 
phology of  this  (or  these)  lizard(s)  is  not  preserved.  This 
material  demonstrates  the  presence  of  at  least  one  other  small 
lizard  in  the  fauna. 

Suborder  Serpentes 
Infraorder  Henophidia 
Family  Boidae 
Subfamily  Erycinae 
Ca/amagras  Cope,  1873 

cf.  Calamagras  sp. 

Figure  3 

MATERIAL.  Centrum  portion  of  trunk  vertebra  retaining 
right  diapophysis  and  portion  of  right  prezygapophysis,  UCR 
21178;  fragment  of  cotyle,  prezygapophysis,  and  diapophy- 
sis, UCR  21179;  centrum  fragment,  UCR  21180. 

DESCRIPTION.  Size  small;  vertebral  centrum  short  (2.35 
mm  from  dorsal  edge  of  cotyle  to  distal  end  of  condyle); 
ventral  surface  of  centrum  smooth  with  no  haemal  keel;  no 
subcentral  ridges;  paradiapophysis  subdivided  into  two  dis- 
tinct articular  surfaces  of  similar  size. 

DISCUSSION.  Although  fragmentary,  this  material  is 


6 Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna 


clearly  referable  to  the  Family  Boidae  on  the  basis  of  the 
short  vertebral  centrum  and  lack  of  haemal  keel.  There  are 
no  neural  arches  or  caudal  vertebrae  necessary  for  positive 
subfamily  assignment,  but  the  small  size  favors  assignment 
to  the  Subfamily  Erycinae.  Among  the  fossil  North  American 
erycines,  only  Calamagras  and  Tregophis  Flolman,  1975, 
completely  lack  a haemal  keel.  The  diagnostic  characters  of 
Tregophis  occur  in  the  neural  arch,  a structure  not  preserved 
in  the  Boron  sample.  Tregophis  is  a peculiar  form  known 
only  from  one  Clarendonian  locality  in  Kansas,  and  thus  it 
appears  more  reasonable  to  refer  the  Boron  species  to  Cala- 
magras, a wide  ranging  taxon  in  the  Middle  Miocene. 

The  cf.  Calamagras  sp.  from  Boron  lacks  the  haemal  keel, 
the  large,  more  widely  separated  paradiapophysial  articular 
surfaces  and  subcentral  gutter  of  Charina  prebottae  Bratt- 
strom,  1958,  from  the  Barstow  Formation. 

The  living  Lichanura  Cope,  1861,  also  lacks  a distinct 
haemal  keel.  The  characters  of  the  neural  spine  and  caudal 
vertebrae  necessary  to  distinguish  Calamagras  from  Licha- 
nura are  not  preserved  in  the  Boron  sample.  Lichanura  has 
not  been  recognized  in  the  pre-Pleistocene  record,  and  the 
Boron  species  is  tentatively  referred  to  Calamagras  for  this 
reason,  not  on  morphological  grounds. 

Family  Colubridae 
Subfamily  Colubrinae 
Paracoluber  Holman,  1970 

cf.  Paracoluber  sp. 

Figure  4 

MATERIAL.  Vertebral  centrum  complete  from  cotyle  to 
condyle  but  lacking  neural  arch  and  associated  structures, 
UCR  21181;  five  uncataloged  posterior  centrum  fragments 
with  condyles. 

DESCRIPTION.  Size  small,  centrum  3.15  mm  long;  cen- 
trum long  compared  to  width;  haemal  keel  moderately  de- 
veloped, oblanceolate,  posterior  end  flat,  not  overlapping 
condyle;  keel  border  straight  in  lateral  view. 

DISCUSSION.  Although  fragmentary,  this  material  is 
complete  enough  to  permit  assignment  to  the  Family  Colu- 
bridae. These  specimens  are  disinguished  from  boids,  the 
common  pre-Barstovian  snakes  in  North  America,  by  the 
long  centrum  and  relatively  narrow,  long  haemal  keel,  from 
the  palaeophids,  elaphids,  viperids  (including  crotalids),  and 
natricine  colubrids  by  the  absence  of  hypapophyses,  and  from 
the  xenodontine  colubrids  (based  on  Heterodon  Latreille, 
1802)  by  the  presence  of  a narrow,  relatively  high  haemal 
keel. 

The  cf.  Paracoluber  sp.  from  Boron  is  distinguished  from 
small  fossil  North  American  colubrines  (see  Holman,  1979) 
by  a combination  of  the  long  centrum,  absence  of  subcentral 
ridges,  and  structure  of  the  haemal  keel.  It  superficially  re- 
sembles Nebraskophis  Holman,  1973,  in  length  of  centrum, 
but  has  a much  deeper  haemal  keel.  The  Boron  species  re- 
sembles Paroxybelis  Auffenberg,  1963,  except  that  the  pos- 


terior end  of  the  haemal  keel  is  not  pointed  and  does  not 
extend  onto  the  condyle.  It  differs  from  Salvadora  pateoli- 
neata  Holman,  1973,  in  lacking  strong  subcentral  ridges. 
Except  for  its  small  size,  cf.  Paracoluber  sp.  from  Boron 
closely  resembles  Paracoluber  storeri  Holman,  1970,  in 
structure  of  the  haemal  keel  and  weak  subcentral  ridges.  Hol- 
man (1970:1322)  indicates  that  a paratype  of  P.  storeri  is 
smaller,  but  does  not  give  the  dimensions. 

The  incomplete  Boron  material  does  not  permit  species 
assignment,  but  it  clearly  demonstrates  the  presence  of  a 
small  colubrid  in  the  fauna. 

Class  Mammalia 
Order  Lagomorpha 

The  lagomorphs  are  the  second  most  common  microverte- 
brates in  the  fauna.  Because  much  of  the  material  is  from 
juveniles,  positive  identification  is  difficult.  Dental  termi- 
nology and  taxonomy  follow  Dawson  (1958). 

Family  Leporidae 
Subfamily  Archaeolaginae 

Archaeolagus  Dice,  1917  or 
Hypo/agus  Dice,  1917 

Archaeolagus  sp.  or  Hypo/agus  sp. 

Figures  5-6 

MATERIAL.  Fragment  of  left  dentary  with  broken  M„ 
complete  M2,  alveolus  for  M3,  UCR  10401;  isolated,  unworn 
LM1,  UCR  10403;  partial  LdP4,  UCR  10400;  isolated,  un- 
worn LP3  or  LdP4,  UCR  10402;  isolated  tooth  and  postcra- 
nial  fragments,  UCR  10404-10412. 

DESCRIPTION.  M'  with  two  lophs  separated  by  cement- 
filled,  lingual  reentrant;  dP4  bilophed  lingually,  enamel  lim- 
ited to  lingual  side,  large,  labially  curved  root  centered  under 
lingual  lophs,  labial  roots  missing;  UCR  10402  (unworn  P3 
or  dP4)  with  distinct  talonid  and  trigonid  separated  by  con- 
tinuous valley  instead  of  separate  lingual  and  labial  reen- 
trants; cement  only  in  central  part  of  valley  separating  talonid 
and  trigonid;  trigonid  composed  of  three  separate  spurs  joined 
at  center  of  tooth,  labial  spur  narrow,  bordered  anterolin- 
gually  by  deep  reentrant. 

DISCUSSION.  Both  the  isolated  M1  and  UCR  10402 
(unworn  P3  or  dP4)  are  from  very  young  individuals,  neither 
showing  wear.  Thus,  both  display  distinct,  separate  cusps  in 
contrast  to  the  lophodont  nature  typical  of  lagomorphs.  The 
bases  of  the  teeth  are  open  and  both  are  relatively  low  crowned, 
suggesting  neither  tooth  was  completely  formed.  The  reen- 
trants of  UCR  10402  (P3  or  dP4)  would  be  persistent  in  wear 
for  nearly  the  entire  length  of  the  preserved  tooth  crown. 
With  some  wear,  the  tooth  would  divide  into  two  lophs  and 
an  additional  anterolabial  reentrant.  Such  a persistent  reen- 
trant is  more  diagnostic  of  Hypo/agus  (Dawson,  1958). 

Archaeolagus  is  typically  characterized  by  the  loss  of  the 
lingual  reentrant  in  the  P3  with  wear,  but  a very  young  in- 


Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna  7 


Figures  3-7.  Stereophotographs  of  cf.  Calamagras  sp.,  cf.  Paracoluber  sp.,  Archaeolagus  sp.  or  Hypolagus  sp.,  and  ? Miospermophilus  sp. 
Figure  3,  cf.  Calamagras  sp.,  UCR  21 178,  ventral  view  of  vertebral  centrum.  Figure  4,  Paracoluber  sp.,  UCR  21181,  ventral  view  of  vertebral 
centrum.  Figures  5-6,  Archaeolagus  sp.  or  Hypolagus  sp.,  UCR  10402,  unworn  LP,  or  dP4;  (5)  occlusal  view,  (6)  labial  view.  Figure  7, 
? Miospermophilus  sp.,  UCR  10399,  unworn  LdP4  or  LP4,  occlusal  view.  Scale  bar  = 1 mm. 


dividual  of  Archaeolagus  acaricolus  Dawson,  1958  (speci- 
men LACM(CIT)  5176)  exhibits  a structure  in  the  P3  similar 
to  UCR  10402.  The  Boron  material  appears  small,  even  for 
Archaeolagus,  but  this  is  probably  due  to  the  juvenile  state 


of  most  of  the  specimens.  The  available  material  from  Boron 
is  not  complete  enough  for  certain  assignment  to  either  Hy- 
polagus or  Archaeolagus.  While  the  size  is  more  consistent 
with  Archaeolagus,  the  morphology  of  UCR  10402,  if  it  is 


8 Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna 


indeed  a P3  in  extremely  early  wear,  more  nearly  resembles 
Hypolagus. 

Order  Rodentia 

The  rodent  fauna  is  dominated  by  heteromyids  except  for  a 
single  small  squirrel  tooth.  Compared  with  other  rodents, 
squirrels  are  generally  uncommon  in  the  fossil  record,  and 
although  the  Boron  species  appears  to  have  some  unique 
characters,  the  single  specimen  is  inadequate  to  make  a spe- 
cific assignment.  The  dental  terminology  used  for  the  squirrel 
follows  Black  (1963). 

At  least  three  heteromyid  species  representing  two  subfam- 
ilies are  present.  As  recognized  by  all  who  work  with  these 
forms,  heteromyid  taxonomy  is  in  great  need  of  revision. 
The  Boron  species  do  not  help  resolve  this  problem,  but  they 
do  indicate  intraspecific  variation  in  some  characters  which 
have  traditionally  been  used  to  separate  species  or  even  gen- 
era. 

The  most  common  species  in  the  microfauna,  represented 
by  more  specimens  than  all  other  microvertebrates  com- 
bined, is  a new  species  of  dipodomyine  heteromyid  referred 
to  Cupidinimus  Wood,  1935.  This  species  represents  the  ear- 
liest record  of  this  widespread  genus  but  it  is  already  clearly 
derived  with  respect  to  species  thought  to  be  ancestral  to  this 
and  closely  related  genera.  The  other  heteromyids,  one  a very 
low  crowned,  bunodont  species  and  the  other  a relatively 
high  crowned  perognathine,  are  each  represented  by  only  two 
teeth  but  are,  nonetheless,  identifiable  at  the  generic  level. 
There  are  several  vials  of  fragmentary  postcranial  bones  from 
small  mammals,  but  none  are  clearly  assignable  to  any  of 
the  species  represented  by  teeth.  Tooth  cusp  terminology  for 
these  heteromyids  is  given  in  Figure  8,  taxonomy  follows 
Wood  (1935)  and  Korth  (1979). 

Family  Sciuridae 
Subfamily  Sciurinae 
Miospermophilus  Black,  1963 

IMiospermoph ilus  sp. 

Figure  7 

MATERIAL.  Unworn,  LdP4  or  LP4,  UCR  10399. 

DESCRIPTION.  Anteroposterior  diameter  = 1.65  mm, 
width  = 1.80  mm;  outline  sub-triangular;  lophs  and  cusps 
high  and  distinct;  metaloph  higher  than  protoloph,  connected 
to  protocone  by  narrow  ridge;  connection  between  protoloph 
and  protcone  stronger  than  connection  between  metaloph 
and  protocone;  protocone  large,  with  separate  posterolingual 
expansion  (possibly  indicating  the  position  of  a separate  hy- 
pocone)  continuous  with  posterior  cingulum;  anterior  and 
posterior  cingula  low  but  distinct;  anterior  cingulum  contin- 
uous from  protocone  to  termination  in  large  parastyle  in 
anterolabial  comer  of  tooth;  posterior  cingulum  running  from 
posterolingual  comer  of  metacone  to  posteriorly  expanded 
protocone  (hypocone);  mesostyle  large. 

DISCUSSION.  The  small  size,  sub-triangular  outline,  rel- 


atively high  cusps  and  lophs  and  presence  of  a metaconule 
serve  to  distinguish  this  species  as  a ground  squirrel  or  chip- 
munk (Black,  1963).  The  Boron  specimen  is  unusual  for 
either  a chipmunk  or  a ground  squirrel  in  the  retention  of 
distinct  cusps,  the  presence  of  a distinct  protoconule,  large 
parastyle,  and  protocone  split  into  two  cusps.  All  these  fea- 
tures might  simply  be  attributable  to  the  unworn  condition 
of  this  specimen,  but  they  also  strongly  suggest  that  it  is  a 
deciduous  premolar.  The  roots  are  missing,  but  the  specimen 
has  the  hollowed  out  base  typical  of  both  deciduous  teeth 
and  unerupted  permanent  teeth. 

Regardless  of  permanent  versus  deciduous  designation,  the 
small  size  favors  assignment  of  the  Boron  specimen  to  one 
of  three  genera,  Miospermophilus  Black,  1963,  Tamias  Illi- 
ger,  1811,  or  Ammospermophilus  Merriam,  1892,  all  rep- 
resented by  Miocene  aged  species.  The  size  is  most  consistent 
with  species  of  Miospermophilus,  which  are  larger  than  those 
of  living  and  extinct  species  of  Tamias  and  the  Miocene 
species  of  Ammospermophilus. 

The  dP4  of  these  small  squirrels  is  poorly  known,  making 
direct  comparison  difficult.  The  dP4  of  M.  bryanti  (Wilson, 
1960)  from  the  Martin  Canyon  Quarry  in  Northeastern  Col- 
orado (in  Black,  1 963: 188)  and  M.  wyomingensis  Black,  1963: 
192  from  the  Split  Rock  Formation  of  Wyoming  are  de- 
scribed as  having  a strong  metaconule  and  low,  distinct  an- 
terior and  posterior  cingula  similar  to  the  Boron  specimen. 
M.  bryanti  lacks  a mesostyle  but  M.  wyomingensis  is  de- 
scribed as  having  a small  mesostyle  similar  to  the  Boron 
specimen.  A partial,  isolated  dP4  from  the  Branch  Canyon 
Formation  of  California  referred  to  Miospermophilus  sp.  by 
Lindsay  (1974: 14)  lacks  the  protoconule  and  metaconule  but 
does  have  a small  mesostyle.  No  described  dP4’s  of  Miosper- 
mophilus have  the  distinct  protoconule  present  in  the  Boron 
specimen,  but  undescribed  specimens  from  the  Thomas  Farm 
faunas  of  Florida  do  have  a protoconule  (D.  Webb  and  A. 
Pratt,  personal  communication). 

The  only  described  dP4  of  a fossil  species  of  Tamias  (Black, 
1963:129)  is  reported  to  bear  the  conules,  large  mesostyle, 
and  expanded  anterior  cingulum  found  in  the  Boron  speci- 
men. 

The  dP4  of  fossil  species  of  Ammospermophilus  has  not 
been  described,  but  specimens  of  this  tooth  and  the  per- 
manent P4  in  a sample  of  the  living  species  A.  lecurus  (Mer- 
riam, 1889)  from  the  LACM  Mammalogy  collections  differ 
considerably  from  the  Boron  specimen.  In  these,  the  anterior 
cingulum  of  the  dP4  extends  only  half  way  along  the  anter- 
olingual  comer  of  the  tooth  and  there  is  no  hint  of  a parastyle 
which  is  prominent  in  the  Boron  IMiospermophilus  speci- 
men. The  metaloph  in  both  the  dP4  and  P4  is  reduced  lin- 
gually  and  does  not  connect  to  the  protocone  even  in  late 
wear  stages  in  the  modem  species  examined.  A further  vari- 
ation was  found  in  specimens  of  the  dP4  of  the  subspecies 
A.  lecurus  peninsulae  { Allen,  1893)  where  the  metaloph  con- 
nects to  the  posterolingual  comer  of  the  lingual  end  of  the 
protoloph. 

Although  similar  in  most  features,  the  permanent  P4  of 
species  of  Miospermophilus,  Tamias,  and  Ammospermophi- 
lus is  generally  even  more  lophodont,  with  greater  reduction 


Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna  9 


hypocone 
hypostyle 

protostyle 


anterior  cingulum  posterior  cingulum 
RM'  Rp‘ 


Figure  8.  Tooth  cusp  terminology  used  in  heteromyid  rodents. 


of  the  metaconule  and  protoconule  than  the  dP4.  The  cingula 
are  also  better  developed  and  nearly  as  high  as  the  protoloph 
and  metaloph  in  the  P4.  The  Boron  specimen  is  similar  in 
size  to  the  P4  of  Miospermophilus  species,  but  differs  in  re- 
tention of  distinct  conules,  particularly  the  protoconule. 

Family  Heteromyidae 
Subfamily  Perognathinae 
Mookomys  Wood,  1931 

Mookomys  sp. 

Figures  9-12,  Table  1 

MATERIAL.  RP4,  UCR  10421;  LM,  or  LM2,  UCR  10464. 

DESCRIPTION.  P4  triangular  in  occlusal  outline  with  four 
subequal  low  rounded  cusps;  protostylid  very  small,  lower 
than  protoconid;  protostylid  and  protoconid  nearly  joined  as 
single  cusp  with  only  a shallow  anterior  groove  separating 
cusps  on  upper  part  of  tooth  crown;  connection  between 
protolophid  and  metalophid  central;  no  posterior  cingulum. 

UCR  10464  (M,  or  M2)  nearly  square  in  occlusal  outline 
with  six  low,  separate  cusps  not  distinctly  connected  until 
late  wear;  tooth  bilophodont  with  wear;  central  valley  deep; 
protostylid  and  hypostylid  small,  both  offset  posteriorly  rel- 
ative to  protoconid  and  hypoconid;  protostylid  anteropos- 
teriorly  expanded;  no  anterior  or  posterior  cingulum;  con- 
nection of  lophids  late,  central;  two  roots. 

DISCUSSION.  The  lower  premolar,  with  its  reduced  pro- 
tolophid, is  more  similar  to  species  of  Heliscomys  Cope, 
1873,  than  to  the  common  species  of  Mookomys,  M.  altiflu- 


Table  1.  Selected  dental  measurements  of  Mookomys  sp.  and  Tro- 
gomys  cf.  T.  rupinimenthae.  Abbreviations:  AP  = anteroposterior  di- 


mension  of  tooth,  WP 
loph(id),  X = mean. 

= width  of  protoloph(id),  WM 

= width  meta- 

UCR 

AP  X WP  X 

WM  X 

Mookomys  sp. 

P4 

10421 

0.70  0.36 

0.73 

M,  or  M2 

10464 

0.86  0.78 

0.79 

Trogomys  cf.  T.  rupinimenthae 

P4 

21182 

1.00 

1.15 

P4 

10419 

0.83  0.60 

0.85 

minis  Wood,  1931,  and  M.  formicorum  Wood,  1935,  both 
with  distinctly  four-cusped  lower  premolars.  The  P4  of  M. 
subtilis  Lindsay,  1972,  is  not  known.  UCR  10421  lacks  the 
posterior  cingulum  diagnostic  of  Heliscomys  species.  A tri- 
angular P4  with  four  cusps,  but  with  a very  small  hypostylid 
and  a narrow  protolophid,  would  be  expected  in  an  early 
species  of  Mookomys  if,  as  is  generally  accepted,  this  genus 
is  a descendant  from  Heliscomys,  which  has  triangular,  three- 
cusped  P4’s. 

The  lower  molar  is  typical  of  species  of  Mookomys  with 
a very  low  crown  and  low  separate  rounded  cusps  which  only 
join  in  later  wear.  The  Boron  specimen  is  much  smaller  than 
that  of  any  species  of  Mookomys  except  Mookomys  subtilis, 
even  if  the  specimen  is  a M2  (which  is  smaller  than  M,).  It 
is  closer  to  size  to  teeth  species  of  Heliscomys,  but  it  lacks 
the  characteristic  cingula.  Mookomys  sp.  from  Boron  differs 
from  the  M , of  M.  subtilis  in  having  a smaller  anterior  cin- 
gulum, less  posteriorly  offset  protostylid  and  hypostylid,  and 
lacking  a posterior  cingulum.  In  this  latter  character  and  the 
very  low  crown  height,  M.  subtilis  is  more  similar  to  species 
of  Heliscomys  than  to  species  of  Mookomys.  Lindsay  (1972) 
does  not  compare  M.  subtilis  (Barstovian)  with  Heliscomys, 
but  it  appears  that  it  is  more  similar  to  Heliscomys,  even 
though  the  latter  is  restricted  to  the  Oligocene. 

Mookomys  sp.  from  Boron  probably  represents  a very  small 
species  of  Mookomys,  but  it  could  be  interpreted  as  a survivor 
species  of  Heliscomys  in  the  early  Hemingfordian. 

Trogomys  Reeder,  1960 
Trogomys  rupinimenthae  Reeder,  1960 

Trogomys  cf.  T.  rupinimenthae 

Figures  13-16,  Table  1 

MATERIAL.  LP4,  UCR  21182;  RP4,  UCR  10419. 

DESCRIPTION.  P4  with  transversely  expanded  proto- 
cone but  no  protostyle;  central  connection  of  protoloph  to 
metaloph;  metaloph  with  three  cusps,  hypocone  and  meta- 
cone closer  together  than  hypocone  and  hypostyle;  hypostyle 
nearly  same  size  as  metacone,  not  anteroposteriorly  expand- 
ed, connected  to  hypocone  by  posterior  cingulum;  three  roots, 
separate  to  base. 


10  Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna 


Figures  9-16.  Stereophotographs  of  Mookomys  sp.  and  Trogomys  cf.  T.  rupinimenthae.  Figures  9-10,  Mookomys  sp.,  UCR  10421,  RP4;  (9) 
occlusal  view,  (10)  lingual  view.  Figures  11-12,  Mookomys  sp.,  UCR  10464,  LM,  or  LM2;  (11)  occlusal  view,  (12)  lingual  view.  Figures  13- 
14,  Trogomys  cf.  T.  rupinimenthae.  UCR  21 182,  LP4;  (13)  labial  view,  (14)  occlusal  view.  Figures  15-16,  Trogomys  cf.  T.  rupinimenthae, 
UCR  10419,  RP4;  (15)  occlusal  view,  (16)  lingual  view.  Scale  bar  = 1 mm. 


P4  four  cusped;  no  hypostylid;  fairly  high  crowned,  clearly 
more  hyposodont  than  Perognathus  Wied-Neuwied,  1839; 
protostylid  smaller  than  protoconid,  offset  posteriorly;  valley 
between  lophids  deep  lingually,  shallow  labially;  first  con- 


nection of  lophids  labial;  sulcus  between  hypoconid  and 
metaconid  as  deep  as  between  metaconid  and  protoconid; 
two  roots,  separate  to  base. 

DISCUSSION.  These  premolars  are  distinctly  lower 


Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna  11 


Figures  17-19.  Stereophotographs  of  Cupidinimus  boronensis  n.  sp.,  UCR  10413,  holotype,  right  dentary  with  dP4,  M,_2;  (17)  occlusal  view, 
(18)  labial  view,  (19)  lingual  view.  Scale  bar  = 2 mm. 


crowned  than  those  of  Cupidinimus,  higher  crowned  and 
more  lophodont  than  those  of  Mookomys  and  somewhat 
higher  crowned  than  those  of  the  common  Barstovian  species, 
Perognathus  furlongi  Gazin,  1930.  The  diagnostic  features 


of  Trogomys,  the  asulcate  upper  incisor  and  characters  of  the 
molars,  are  not  represented  in  the  Boron  sample.  The  two 
Boron  teeth  are  within  the  size  range  of  P.  furlongi,  but  are 
higher  crowned,  a characteristic  of  Trogomys.  They  are  com- 


12  Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna 


Table  2.  Selected  dental  measurements  of  Cupidinimus  boronensis 
n.  sp.  Abbreviations:  AP  = anteroposterior  dimension  of  tooth,  WP 
= width  protoloph(id),  WM  = width  metaloph(id),  X = mean. 


UCR 

AP 

X 

WP 

X 

WM 

X 

Type  M, 

10413 

0.98 

1.19 

1.30 

m2 

0.91 

1.15 

1.16 

dP4 

1.42 

0.97 

P4 

10449 

1.30 

1.26 

1.40 

1.31 

10450 

1.31 

1.27 

10451 

1.30 

1.25 

10452 

1.12 

1.31 

10453 

1.20 

1.29 

10454 

1.45 

1.37 

10456 

1.23 

1.36 

10457 

1.13 

1.29 

10458 

1.26 

1.29 

M1 

10459 

0.95 

0.99 

1.37 

1.30 

1.23 

1.16 

10460 

0.95 

1.17 

1.10 

10461 

1.03 

1.36 

1.25 

10462 

1.00 

1.34 

1.20 

10463 

1.02 

1.28 

1.16 

21183 

1.00 

1.25 

1.00 

M2 

10465 

0.90 

0.86 

1.23 

1.11 

1.02 

0.99 

10467 

0.88 

1.04 

0.92 

10468 

0.83 

1.11 

1.04 

21184 

0.83 

1.05 

0.98 

M3 

10466 

0.77 

0.87 

- 

dPJ 

10471 

1.08 

1.01 

— 

10472 

1.19 

1.15 

- 

P4 

10416 

0.95 

1.00 

0.91 

0.88 

1.05 

1.05 

10417 

1.11 

0.89 

1.13 

10418 

1.02 

0.83 

1.10 

10420 

0.99 

0.90 

0.95 

10422 

0.94 

0.88 

1.03 

M, 

10413 

0.98 

1.05 

1.19 

1.17 

1.30 

1.27 

10423 

1.04 

1.16 

1.29 

10424 

1.06 

1.29 

1.29 

10427 

1.09 

1.15 

1.27 

10429 

1.02 

1.23 

1.27 

10434 

1.15 

1.19 

1.28 

10436 

1.08 

1.19 

1.27 

10438 

1.06 

1.13 

1.19 

10439 

1.00 

1.13 

1.25 

10441 

1.09 

1.05 

1.22 

M, 

10413 

0.91 

0.95 

1.15 

1.19 

1.16 

1.10 

10426 

0.90 

1.17 

1.08 

10430 

1.03 

1.21 

1.09 

10431 

0.89 

1.25 

1.11 

10432 

0.90 

1.19 

— 

10433 

1.05 

1.23 

1.10 

10435 

1.01 

1.24 

1.07 

Table  2.  Continued. 


UCR 

AP 

X 

WP 

X 

WM 

X 

10437 

0.87 

1.13 

1.15 

10440 

1.04 

1.23 

1.16 

10444 

0.96 

1.26 

1.07 

10447 

0.88 

1.05 

1.00 

M, 

10443 

0.74 

0.73 

1.08 

0.92 

0.77 

0.76 

10445 

0.74 

0.91 

0.76 

10446 

0.72 

0.76 

0.74 

dP4 

10414 

1.32 

- 

0.75 

parable  in  size,  crown  height,  and  cusp  morphology  to  T. 
rupinimenthae  from  the  Arikareean  age  Tick  Canyon  For- 
mation of  southern  California. 

Subfamily  Dipodomyinae 
Cupidinimus  Wood,  1935 

Cupidinimus  Wood,  1935:1  18. 

Perognathoides  W ood,  1935:92. 

Prodipodomysl  mascallensis  Downs,  1956;  sample  of  Shot- 
well,  1967:22. 

Perognathus  saskatchewanensis  Storer,  1 970: 1127. 

Cupidinimus  boronensis  new  species 

Figures  17-39,  Table  2 

HOLOTYPE.  UCR  10413,  right  dentary  with  incisor,  dP4, 

M,_2. 

TYPE  LOCALITY.  UCR  locality  RV  642 1 , Arkose  Mem- 
ber of  the  Kramer  Beds  of  Barnard  and  Kistler  (1966),  146 
m above  the  top  of  the  Saddleback  Basalt  on  the  third  level 
below  the  surface  in  the  east  wall  of  the  open  pit  mine  of  the 
United  States  Borax  and  Chemical  Corporation  as  it  existed 
in  March,  1964,  4.8  km  northwest  of  Boron,  Section  23, 
T.  1 IN.,  R.8W.,  Boron  Quadrangle,  United  States  Geological 
Survey,  1954,  1:62,500,  San  Bernardino  County,  California. 

REFERRED  MATERIAL.  Five  RP4’s,  UCR  10449,  UCR 
10450,  UCR  10451, UCR  10452, and  UCR  10453;five  LP4’s, 
UCR  10454,  UCR  10455,  UCR  10456,  UCR  10457,  and 
UCR  10458;  two  RM"s,  UCR  10460  and  UCR  21183;  four 
LM"s,  UCR  10459,  UCR  10461,  UCR  10462,  and  UCR 
10463;  two  RM2’s,  UCR  10465  and  UCR  21 184;  two  LM2’s, 
UCR  10467  and  UCR  10468;  RM3,  UCR  10466;  LM1  or 
LM2,  UCR  1 0469;  two  RdP4’s,  UCR  10471  and  UCR  10472; 
two  RP4’s,  UCR  10418  and  UCR  10420;  four  LP4’s,  UCR 
10415,  UCR  10416,  UCR  10417,  and  UCR  10422;  seven 
RM.’s,  UCR  10423,  UCR  10424,  UCR  10427,  UCR  10436, 
UCR  10438,  UCR  10439,  and  UCR  10441;  four  LM.’s, 
UCR  10428,  UCR  10429,  UCR  10434,  and  UCR  10470; 
five  RM2’s,  UCR  10431,  UCR  10437,  UCR  10440,  UCR 
10444,  and  UCR  10447;  six  LM,’s,  UCR  10425,  UCR  10430, 
UCR  10432,  UCR  10433,  UCR  10435,  and  UCR  10442; 
RM3,  UCR  10445;  two  LM3’s,  UCR  10443  and  UCR  10446; 


Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna  13 


Figures  20-29.  Stereophotographs  of  Cupidinimus  boronensis  n.  sp.  lower  dentition.  Figure  20,  UCR  10413,  holotype,  occlusal  view  of 
dentition  only.  Figues  21-22,  UCR  10416,  LP4;  (21)  occlusal  view,  (22)  labial  view.  Figure  23,  UCR  10422,  LP4,  occlusal  view.  Figures  24- 
25,  UCR  10423,  RM,;  (24)  occlusal  view,  (25)  labial  view.  Figures  26-27,  UCR  10430,  LM2;  (26)  occlusal  view,  (27)  lingual  view.  Figure  28, 
UCR  10445,  RM,,  occlusal  view.  Figure  29,  UCR  10414,  RdP4,  occlusal  view.  Scale  bar  = 1 mm. 


14  Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna 


RdP4,  UCR  10414;  all  from  type  locality,  UCR  locality  RV 
6421. 

DIAGNOSIS.  Teeth  typical  of  dipodomyines  by  being 
higher  crowned  than  those  of  perognathines  and  most  het- 
eromyines;  further  distinguished  from  heteromyines  by  lack- 
ing a “J”  wear  pattern  in  the  P4;  distinguished  from  geo- 
myines  by  smaller  size,  lower  crown  height,  retention  of 
cuspidate  teeth  and  central  (versus  lingual)  connection  of 
lophs  in  P4;  individual  teeth  larger  than  those  of  C.  nebras- 
kensis  Wood,  1935,  C.  halli  (Wood,  1936b),  C.  cuyamensis 
(Wood,  1937),  and  C.  madisonensis  (Dorr,  1956),  smaller 
than  C.  bidahochiensis  (Baskin,  1979);  teeth  high  crowned 
but  retaining  individual  cusps  until  moderate  wear;  roots 
long,  separate  to  base;  premolars  lacking  accessory  cusp(id)s; 
M33  reduced;  dP44  low  crowned,  with  complex  cusp  pattern; 
metaloph  of  P4  wide  compared  to  width  of  M1,  hypostyle 
not  greatly  elongate  and  posterior  cingulum  incomplete;  P4 
short  anteroposteriorly  relative  to  width  of  protolophid  and 
length  of  M,;  anterior  cingulum  not  anteriorly  expanded  in 
M,  or  M2;  dP4  with  large  anteroconid  connected  to  central 
mure  and  lacking  hypostylid;  in  addition  to  size,  differs  from 
C.  halli  and  C.  madisonensis  by  lack  of  accessory  cusps  in 
P44,  anteroposteriorly  short  P4,  reduced  anterior  cingula  on 
M,  and  M2  and  large  anteroconid  connected  to  central  mure 
and  lack  of  hypostylid  in  dP4;  in  addition  to  size,  differs  from 
C.  nebraskensis  by  anteroposteriorly  short  P4,  relatively  un- 
reduced M2,  reduced  M33,  and  strong  central  mure  on  dP4. 

ETYMOLOGY.  Named  after  the  type  locality. 

DESCRIPTION.  P4  with  transversely  expanded  proto- 
cone; accessory  cusps  lacking  on  protocone  except  for  one 
specimen,  UCR  10454,  with  greater  transverse  expansion  of 
protoloph  suggestive  of  protostyle;  first  connection  of  lophs 
central;  metaloph  three  cusped;  hypocone  slightly  posterior 
relative  to  lateral  cusps;  hypostyle  slightly  expanded  antero- 
posteriorly; posterior  cingulum  small,  between  postero- 
medial surface  of  hypocone  and  hypostyle;  three  roots,  one 
each  under  protoloph,  metacone,  and  hypostyle,  separate  to 
base. 

M1  six  cusped  in  early  wear,  cusps  less  persistent  than  in 
lower  molars;  anterior  cingulum  low  relative  to  protocone, 
continuous  between  paracone  and  protostyle,  thickening  near 
paracone;  protoloph  wider  than  metaloph;  anterior  face  of 
protoloph  flat;  small  protolophule  connecting  protocone  and 
paracone  in  2 of  3 specimens  with  early  enough  wear  to  show 
character;  transverse  valley  deep  and  narrow;  first  union  of 
lophs  lingual,  followed  by  central  connection,  but  no  lake 
formation;  median  cusps  (protocone  and  hypocone)  posi- 
tioned posteriorly  relative  to  lateral  cusps,  making  metaloph 
convex  posteriorly;  no  posterior  cingulum  except  for  con- 
nection between  hypocone  and  hypostyle;  one  lingual,  two 
labial  roots,  separate  to  base. 

M2  smaller  than  M\  proportionately  more  reduced  than 
M2;  anterior  face  convex;  posterior  face  concave;  anterior 
cingulum  reduced  to  absent;  protolophule  between  protocone 


and  paracone  present  in  one  specimen;  first  union  of  lophs 
labial,  but  transverse  valley  shallower  than  in  M 1 , closed  both 
lingually  and  labially;  metaloph  narrower  than  protoloph  due 
to  reduction  in  metacone;  metaloph  three  cusped,  joined  into 
continuous  posterior  crescent;  hypostyle  offset  relative  to  hy- 
pocone and  more  anteroposteriorly  elongate  than  in  M1;  one 
lingual,  two  labial  roots,  separate  to  base. 

M3  with  no  anterior  cingulum;  protostyle  elongate  antero- 
posteriorly; metaloph  bearing  three  cusps  joined  to  posterior 
crescent  which  lacks  distinct  cusps;  roots  not  preserved  in 
available  sample. 

dP4  low  crowned;  triangular  outline;  cusps  low  but  distinct; 
three  lophed;  large,  transversely  expanded  anterocone;  pro- 
toloph with  two  cusps,  larger  protocone  and  smaller  para- 
cone; metaloph  with  three  cusps,  metacone,  hypocone,  and 
anteroposteriorly  expanded  hypostyle;  anterolabial  connec- 
tion between  protoloph  and  anterocone;  no  connection  be- 
tween protoloph  and  metaloph;  hypostyle  extends  anteriorly 
to  lingual  side  of  protocone  but  with  no  connection  to  pro- 
tocone; three  roots,  one  each  under  anterocone,  metacone, 
and  protostyle. 

P4  with  four  cusps  nearly  equal  in  size;  protostylid  smaller 
than  protoconid,  closer  to  hypoconid  than  protoconid  is  to 
metaconid;  protoconid-protostylid  connection  central;  pro- 
tolophid narrower  than  metalophid;  first  connection  of  lophs 
either  slightly  lingual  of  center,  between  protostylid  and  hy- 
poconid, or  central;  two  roots,  separate  to  base. 

M,  six  cusped;  tooth  wider  than  long;  protostylid  nearly 
same  size  as  protoconid,  offset  posteriorly  relative  to  pro- 
toconid, continuous  with  anterior  cingulum  which  connects 
to  anteromedial  surface  of  protoconid;  metalophid  convex 
anteriorly,  hypolophid  straight;  transverse  valley  deep,  nar- 
row, open  labially  and  lingually,  deeper  on  lingual  side;  first 
union  of  lophs  both  between  protostylid  and  hypostylid  and 
central,  probably  forming  short-lived  lake;  hypolophid  nar- 
rower than  metalophid  due  to  expanded  entoconid;  hypo- 
stylid smaller  than  hypoconid;  no  posterior  cingulum;  two 
roots,  separate  to  base. 

M2  six  cusped;  tooth  wider  than  long;  metalophid  and 
hypolophid  same  width;  metalophid  less  convex  anteriorly 
than  in  M,;  protostylid  small,  smaller  than  in  M„  offset 
posteriorly  relative  to  protoconid,  continuous  with  anterior 
cingulum  which  extends  only  to  anterolabial  corner  of  pro- 
toconid; hypostylid  smaller  than  on  M,;  transverse  valley 
deep,  deeper  lingually  than  labially,  labial  connection  deeper 
than  in  M,;  first  union  of  lophs  more  strongly  central  than 
in  M„  labial  connection  late  or  lacking;  hypolophid  straight; 
two  roots,  separate  to  base;  two  early  wear  specimens,  UCR 
1 0442  and  1 0444  displaying  variation  from  above  by  lacking 
distinct  hypostylid  (but  with  hypoconid  extending  labially 
toward  base,  thus  forming  as  broad  a hypolophid  as  an  M2 
with  hypostylid)  and  with  protostylid  separated  from  anterior 
cingulum  by  distinct,  shallow  groove  lost  with  early  wear. 

M3  four  cusped,  lacking  stylids;  bilophodont  in  early  wear; 


Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna  15 


4 

30 

32 

•1 

tv 

31 

n 11 

33 

n 

0 

35 

0 0 

34 

© 

36 

W 6 

38 

w 

37 

39 

Figures  30-39.  Stereophotographs  of  Cupidinimus  boronensis  n.  sp.  upper  dentition.  Figures  30-31,  UCR  10450,  RP4;  (30)  occlusal  view, 
(31)  labial  view.  Figure  32,  UCR  10449,  RPJ,  occlusal  view.  Figures  33-34,  UCR  10460,  RM1;  (33)  labial  view,  (34)  occlusal  view.  Figure 
35,  UCR  10467,  LM2,  occlusal  view.  Figure  36,  UCR  10445,  RM3,  occlusal  view.  Figure  37,  UCR  10472,  RdP\  occlusal  view.  Figures  38- 
39,  UCR  10471,  RdPJ;  (38)  lingual  view,  (39)  occlusal  view.  Scale  bar  = 1 mm. 


16  Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna 


very  small  anterior  cingulum  in  early  wear,  forming  anter- 
olabial  “comer”  on  protoconid  with  wear;  metalophid  dis- 
tinctly wider  than  hypolophid;  central  valley  narrow,  deep; 
single  rooted. 

dP4  very  low  crowned;  elongate  oval  in  occlusal  view,  nar- 
rower anteriorly;  complex  cusp  pattern  with  three  main  cen- 
tral cusps,  metaconid,  entoconid,  and  hypoconid;  large  an- 
terolingually  placed  anteroconid  nearly  continuous  with  high 
anterolabial  cingulum;  anterolabial  cingulum  and  protostylid 
as  high  as  central  cusps,  nearly  continuous  to  mid-labial  por- 
tion of  tooth  and  connecting  back  lingually  to  central  mure; 
very  small  cuspule  between  hypoconid  and  posterior  end  of 
anterolabial  cingulum;  low  protoconid  anterolabial  to  meta- 
conid; no  hypostylid;  with  wear,  hypoconid  joins  a central 
mure  which  runs  anteriorly  through  protoconid  to  antero- 
conid; with  wear,  metaconid  joined  by  short  loph  to  proto- 
conid; entoconid  connected  by  mure  either  directly  to  hy- 
poconid or  through  posterior  cingulum  to  hypoconid;  small 
lingual  accessory  cuspid  between  metaconid  and  entoconid; 
large  posterior  cingulum;  two  roots,  one  anterior,  one  pos- 
terior. 

DISCUSSION.  C.  boronensis  is  assigned  to  the  genus 
Cupidinimus  following  the  expanded  concept  of  this  genus 
proposed  by  Korth  (1979).  As  previously  noted  by  Reeder 
(1956),  Lindsay  (1972),  and  Sutton  (1977),  the  genus  Cupi- 
dinimus, represented  by  the  single  species  C.  nebraskensis, 
and  at  least  some  of  the  eight  species  referred  to  Perogna- 
thoides,  are  nearly  indistinguishable  in  dental  morphology. 
Characters  which  have  been  used  to  distinguish  these  species 
were  found  to  be  variable  as  larger  samples  became  available. 
Thus,  Korth  (1979)  proposed  formal  synonymy  of  Perog- 
nathoides  and  several  other  species  referred  to  other  genera 
with  Cupidinimus. 

C.  boronensis  is  typical  of  this  expanded  Cupidinimus 
species  complex  with  high-crowned  teeth  which  retain  in- 
dividual cusps  until  moderate  wear.  It  is  unique  in  combining 
large  size,  length  (anteroposterior)  of  P4  much  less  than  M, 
(a  Perognathus- like  character),  high-crowned  teeth,  relatively 
unreduced  M2,  reduced  M33,  and  dP4  with  central  mure  and 
no  hypostylid  (C.  halli-C.  madisonensis  characters),  with  a 
lack  of  accessory  cusps  on  P44,  a wide  metaloph  on  P4  com- 
pared to  M1,  a reduced  anterior  cingulum  on  M,,  and  dP4 
with  large  anteroconid  connected  to  central  mure  (C.  ne- 
braskensis characters). 

C.  boronensis  is  higher  crowned  than  the  perognathines 
Mookomys,  Perognathus,  and  Trogomys.  It  is  also  higher 
crowned  than  most  species  of  the  heteromyines  Peridiomys 
Matthew,  1924,  and  Proheteromys  Wood,  1932.  These  and 
the  higher  crowned  heteromyine  Diprionomys  Kellogg,  1910, 
are  further  distinguished  by  the  “J”  wear  pattern  in  the  P4 
with  the  first  union  of  lophs  lingually  between  the  protocone 
and  hypostyle.  C.  boronensis  is  smaller,  lower  crowned,  and 
lacks  the  labial  connection  of  protoloph  and  metaloph  in  the 
P4  of  Mojavemys  Lindsay,  1972,  and  other  geomyines. 


C.  boronensis  is  closest  in  size  to  C.  halli  and  C.  madi- 
sonensis (=  C.  halli  according  to  Lindsay,  1972,  but  consid- 
ered distinct  by  Sutton,  1 977,  a view  I will  follow).  The  molars 
are  as  large  as  or  larger  than  those  of  C.  halli  and  C.  madi- 
sonensis. C.  boronensis  also  differs  from  these  species  as  fol- 
lows: 1)  the  roots  are  less  fused,  2)  the  length  of  P4  is  less  in 
comparison  to  the  width  of  the  metalophid  and  length  of  M,, 
3)  the  P4  lacks  an  anteroconid,  and  the  connection  of  pro- 
toconid and  protostylid  is  central  rather  than  posterior,  4) 
the  relative  shortness  of  the  P4  is  similar  to  the  condition  in 
Perognathus  and  Trogomys,  in  contrast  to  other  species  of 
Cupidinimus  in  which  this  is  the  longest  tooth  in  the  lower 
tooth  row,  5)  the  anterior  cingulum  of  C.  boronensis  is  small- 
er and  less  angular  in  both  M,  and  M2,  connecting  only  to 
the  protostylid,  not  the  protostylid  and  metaconid,  6)  the 
hypostylid  is  lacking  in  2 of  12  specimens  of  M„  7)  the  P4 
lacks  accessory  cusps  on  the  protocone,  the  hypocone  is  less 
posterior  relative  to  lateral  cusps,  the  hypostyle  and  posterior 
cingulum  are  smaller,  and  the  metalophid  is  wider  relative 
to  the  width  of  M1,  8)  the  M1  lacks  a posterior  cingulum  and 
the  protolophule  between  protocone  and  paracone  is  smaller, 
9)  compared  to  specimens  referred  to  C.  halli  by  Lindsay 
(1972),  the  dP4  is  much  lower  crowned  with  a larger  anter- 
olabial cingulum,  smaller  anteroconid,  no  hypostylid,  and  a 
weaker  central  mure  (except  that  it  is  fully  connected  to  an- 
teroconid), and  10)  the  dP4,  even  though  unworn,  is  lower 
crowned,  lacks  a connection  between  hypostyle  and  proto- 
cone and  has  a labial  connection  between  the  protoloph  and 
anterocone. 

C.  boronensis  is  1 8-20  percent  larger  than  C.  nebraskensis 
from  the  type  area  (Korth,  1979)  and  the  sample  from  the 
Barstow  Formation  (Lindsay,  1972).  It  resembles  the  sample 
from  the  Barstow  Formation  in  lacking  the  anteroconid  on 
P4,  but  this  character  is  variably  developed  in  the  topotypic 
sample  from  the  Valentine  Formation.  C.  boronensis  lacks 
the  relative  reduction  of  the  M2  found  in  C.  nebraskensis  but 
has  a proportionally  more  reduced  M3. 

C.  boronensis  differs  from  species  of  Cupidinimus  previ- 
ously referred  to  Perognathoides,  C.  cuyamensis,  C.  klein- 
felderi  (Storer,  1970),  and  C.  bidahochiensis,  by  lacking  ac- 
cessory cusps  in  the  premolars  variously  developed  in  all 
these  and  differs  further  from  C.  bidahochiensis  by  much 
smaller  size.  C.  boronensis  also  differs  from  C.  kleinfelderi, 
known  only  from  isolated  premolars,  by  lacking  an  anteriorly 
drawn  out  protolph(id)  compared  to  metaloph(id)  on  P44. 

The  types  of  C.  eurekensis  (Lindsay,  1972),  C.  quartus 
(Hall,  1930),  and  C.  tertius  (Hall,  1930),  all  previously  re- 
ferred to  Perognathoides,  have  highly  worn  teeth  which  lack 
diagnostic  characters  that  would  facilitate  comparison  with 
C.  boronensis,  but  C.  eurekensis  is  considerably  smaller  and 
C.  quartus  and  C.  tertius  are  larger. 

C.  boronensis  differs  from  four  other  species  referred  to 
Cupidinimus  by  Korth  (1979),  a sample  of  C.  cf.  C.  tertius 
from  the  Avawatz  Formation  (Wilson,  1939),  C.  cf.  C.  cuya- 


Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna  17 


Table  3.  Selected  measurements  of  Merychyus  cf.  M.  minimus  and 
IMerychyus  sp.  from  the  Boron  Local  Fauna.  Abbreviations:  AP  = 
anteroposterior  dimension  of  tooth,  TR  = width  of  tooth  at  anterior 
loph(id),  CH  = crown  height  at  mesostyle  in  uppers,  at  metaconid 
in  lowers,  * = approximate. 


Skull 

Merychyus  cf.  M.  minimus 

IMery- 
chyus  sp. 

UCR  UCR 

10319  10320 

UCR 

10321 

UCR 

10322 

UCR 

10331 

Facial  length,  C to 

anterior  rim  orbit 

52.8 

51.1 

54* 

Jugal  depth  below  orbit 

13.2 

12.8 

17.7 

Length  P'-P4 

30.0 

30.8 

33* 

Length  M'-M3 

38.0 

44.7 

Length  C-M3 

74.3 

- 

P2  AP 

7.8 

8.0 

7.8 

TR 

6.0 

6.8 

6.6 

P3  AP 

8.1 

9.6 

8.2 

9.2 

TR 

7.6 

8.0 

7.1 

7.5 

P4  AP 

7.4 

7.3 

7.2 

TR 

9.2 

8.6 

10.1 

CH 

10.4 

7.6 

M1  AP 

— 

11.0 

12.8 

1 1.5 

TR 

— 

11.1 

10.8 

1 1* 

CH 

— 

9.1 

10.5 

- 

M2  AP 

13.5 

15.4 

— 

13.4 

TR 

1 1.0 

12.5* 

13.8 

CH 

— 

12.6 

— 

- 

M3  AP 

18.0 

21.0 

TR 

13.0 

14.0 

CH 

14.7 

8.9 

Length  dP2-dP4 

25.7 

26.8 

Length  dP2 

7.4 

Length  dP3 

8.7 

10.1 

Length  dP4 

8.7 

9.4 

Merychyus  cf. 

M.  minimus 

1 Merychyus  sp. 

UCR 

UCR 

UCR 

UCR 

Dentary 

10323  1 

10325 

10332 

21270 

Mandible  depth  below  P2 

22.6 

17.1 

24.5 

Length  of  symphysis 

33.8 

22.0 

32.3 

Length  C-M, 

79.8 

80.1 

Length  P,-P4 

31.9 

29.5 

Length  M,-M3 

44.2 

47.0 

M,  AP 

12.6 

12.9 

TR 

8.0 

8.2 

CH 

5.5 

8.7 

M,  AP 

19.8 

23* 

21.5 

TR 

8.9 

8.8 

8.9 

CH 

— 

17.0 

Length  dP3-dP4 

31.8 

mensis  from  the  Valentine  Formation  (Klinginger,  1968),  a 
sample  referred  to  as  “ Prodipodomys"  ? mascallensis  (Downs, 
1956),  from  the  Quartz  Basin  in  Oregon  (in  Shotwell,  1967), 
and  C.  saskatchewanensis  (Storer,  1975),  by  lacking  acces- 
sory cusps  on  the  premolars.  C.  boronensis  is  close  in  size  to 
the  Quartz  Basin  C.  mascallensis  and  otherwise  similar  ex- 
cept that  the  connection  of  the  protoloph  and  metaloph  in 
P4  is  lingual  rather  than  central  in  C.  mascallensis.  C.  bo- 
ronensis is  also  similar  in  size  to  C.  saskatchewanensis,  but 
the  latter  is  lower  crowned  and  the  protolophid  of  its  P4  is 
relatively  anteriorly  drawn  out  as  in  most  species  of  Cupi- 
dinimus. 

C.  boronensis  is  conservative  among  species  of  Cupidini- 
mus  in  lacking  development  of  accessory  cusps  on  the  pre- 
molars. It  is  even  larger  than  Barstovian  species,  in  contrast 
to  an  apparent  general  trend  of  size  increase  from  Barstovian 
to  Hemphillian.  Although  larger  than  the  common  Barsto- 
vian C.  halli,  it  is  lower  crowned,  more  like  the  much  smaller, 
contemporaneous  and  longer  surviving  C.  nebraskensis.  This 
combination  of  characters  would  suggest  that  C.  boronensis 
is  not  directly  ancestral  to  either  the  smaller,  higher  crowned 
C.  halli  or  the  smaller  C.  nebraskensis.  It  may  represent  a 
separate  lineage  that  increased  in  size  early  or,  by  reversal 
of  the  size  increase,  gave  rise  to  C.  nebraskensis  or  similar 
species. 

C.  boronensis  could  easily  be  derived  from  any  perogna- 
thine  by  a significant  increase  in  crown  height  and  a reduction 
of  cingula.  Trogomys  was  suggested  as  a likely  ancestor  to 
the  C.  halli/C.  nebraskensis  complex  in  the  Barstow  For- 
mation by  Lindsay  (1972).  C.  boronensis  is  much  higher 
crowned  and  has  much  reduced  cingula  compared  to  the  type 
of  Trogomys,  and  it  is  only  separated  from  the  latter  by  a 
short  geologic  interval.  However,  Trogomys  is  the  most  likely 
candidate  among  the  perognathines. 

C.  boronensis  is  the  earliest  species  in  the  Cupidinimus 
species  group  and  the  only  Hemingfordian  representative  of 
this  successful  group.  In  general,  Hemingfordian  heteromyids 
are  poorly  known,  and  as  such,  C.  boronensis  provides  a rare 
look  at  these  small  rodents  in  the  time  span  between  Ari- 
kareean  faunas  dominated  by  pleurolicine,  primitive-pe- 
rognathine,  and  Proheteromys  spp.  heteromyid  assemblages 
and  Barstovian  faunas  dominated  by  Cupidinimus  spp.  and 
advanced-perognathine  heteromyid  assemblages. 

Order  Artiodactyla 

The  artiodactyls  are  represented  by  three  families,  oreodonts, 
camels,  and  palaeomerycids.  More  than  half  the  specimens 
are  juveniles,  making  positive  identifications  difficult  in  some 
cases. 

The  oreodonts,  represented  by  one  or  more  small,  sub- 
hypsodont  species,  are  more  common  than  the  camels.  Ore- 
odont  taxonomy  is  in  need  of  revision  and  the  material  from 
Boron  sheds  little  light  on  this  vexing  problem.  Taxonomy 
used  follows  Schultz  and  Falkenbach  (1947)  and  Stevens 
(1977). 

As  is  typical  of  Miocene  faunas,  the  Boron  Local  Fauna 
contains  at  least  three  camel  taxa.  Most  of  the  material  is 


18  Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna 


Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna  19 


Figures  40-43.  Merychyus  cf.  M.  minimus.  Figures  40-41,  UCR  10319,  partial  cranium  with  right  facial  region,  RI1-3  (roots  only)  and  RC-RM3;  (40)  lateral  view,  (41)  occlusal 
view.  Figures  42-43,  UCR  10323,  lower  mandible  with  complete  dentition;  (42)  left  lateral  view,  (43)  occlusal  view.  All  actual  size. 


I 


Figures  44-45.  Merychyus  cf.  M.  minimus,  UCR  10320,  juvenile  cranium  with  C,  P1,  LdP3-4,  M1-2,  unerupted  RP2-4  exposed  by  removing 
deciduous  right  premolars;  (44)  dorsal  view,  (45)  occlusal  view.  All  actual  size. 

poorly  preserved.  Only  two  species  are  represented  by  partial  Webb  (1965),  and  Honey  and  Taylor  (1978);  osteological 
dentitions  (one  only  by  juvenile  dentition)  and  associations  terminology  follows  Webb  (1965). 

with  postcranial  elements  have  been  made  on  size  alone.  The  unique  artiodactyl  in  the  Boron  Local  Fauna  is  a new 

Thus,  only  one  camel  is  tentatively  identified  at  the  generic  species  of  hypsobrachyodont  aletomerycine  palaeomerycid. 

level.  The  other  two  cannot  even  be  confidently  assigned  to  Whereas  palaeomerycids  are  relatively  uncommon  in  fossil 

subfamily  level.  The  taxonomy  follows  Macdonald  (1949),  assemblages  of  the  Great  Basin  and  southern  California  when 


20  Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna 


compared  to  other  artiodactyls,  this  species  is  the  most  abun- 
dant large  animal  at  Boron  and  is  exceeded  in  numbers  of 
specimens  only  by  Cupidinimus  boronensis.  Palaeomerycid 
taxonomy  is  traditionally  based  on  the  orbital  horns  common 
in  this  group.  The  new  species  from  Boron  is  represented  by 
one  horn  fragment,  but,  in  this  case,  the  more  diagnostic 
characters  are  in  the  dentition.  This  new  species  is  notewor- 
thy because  it  is  the  first  well-documented  aletomerycine 
found  west  of  the  Rocky  Mountains,  thus  demonstrating  a 
broader  distribution  than  previously  recognized  for  these 
small,  hypsobrachyodont  “homed  ruminants.”  Dental  ter- 
minology used  is  given  in  Figure  50  and  taxonomy  follows 
Frick  (1937). 

Family  Merycoidodontidae 
Subfamily  Merychyinae 
Merychyus  Leidy,  1858 
Merychyus  minimus  Peterson,  1906 

Merychyus  cf.  M.  minimus 

Figures  40-45,  Table  3 

MATERIAL.  Fragment  of  cranium  with  right  orbit  and 
facial  region,  RI1'3  (roots  only),  and  RC-M\  UCR  10319; 
juvenile  cranium  lacking  posterior  portion  behind  orbits,  with 
moderately  worn  C,  P1,  dP3~4,  M‘~2,  and  unerupted  P2-4  ex- 
posed on  right  side,  UCR  10320;  juvenile  palate  with  well 
worn  dl1-3,  dC,  P1,  dP2~4,  and  slightly  worn  M1-2,  UCR  10321; 
partial  left  maxilla  with  moderately  worn  I3,  C,  P1-3,  UCR 
10322;  isolated  LM3,  UCR  21185;  mandibular  fragment  with 
complete  dentition,  lacking  ascending  ramus,  angle  and  con- 
dyles, UCR  10323;  natural  mold  of  right  dentary  with  con- 
dyle, P4-M3,  UCR  10324;  juvenile  mandible  with  partially 
erupted  P,  slightly  worn  dC,  dP,_3,  and  M,,  UCR  10325; 
unworn  LM,  in  maxilla  fragment,  UCR  10326. 

DESCRIPTION.  Size  (based  on  length  of  upper  tooth  row) 
slightly  larger  than  average  Merychyus  crabilli  Schultz  and 
Falkenbach,  1947,  and  Merychyus  calaminthus  Jahns,  1940, 
within  size  range  of  average  Merychyus  minimus;  skull  with 
shallow  antorbital  fossa  and  narrow  lacrimal  vacuity;  jugal 
shallow;  orbit  large;  two  infraorbital  foramina  with  posterior 
one  above  P4;  dentition  sub-hypsodont  (ratio  of  height  of 
enamel  of  paracone  of  M3  to  width  of  M3  = 0.9);  molars 
appear  narrow  in  relation  to  length  even  in  late  wear  giving 
teeth  less  robust  appearance  than  typical  in  oreodonts;  pre- 
molars large;  ratio  of  length  of  P1-4  to  length  of  M1-3  = 0.81 
(premolar  diameter  of  Stevens,  1977);  premolars  lacking 
complexity  of  spurs  typical  in  Merychyus  ( Metoreodon ) re- 
lictus  Matthew  and  Cook,  1 909;  upper  premolar  spacing  closed 
but  not  crowded;  lower  premolar  spacing  somewhat  open; 
anterior  margins  of  P 1-3  slant  backward;  upper  C and  P,  not 
enlarged. 

DISCUSSION.  Historically,  oreodont  taxonomy  has  been 
based  on  samples  of  complete  skulls.  The  fragmentary  ma- 
terial from  Boron  does  not  permit  such  an  analysis,  and  the 
study  is  further  complicated  by  the  large  amount  of  juvenile 
material.  Oreodonts  appear  to  be  evolutionarily  conservative 


in  dental  characters.  Even  generic  and  subfamily  differences 
are  difficult  to  distinguish  using  only  the  dentition  and  there 
is  little  agreement  among  workers  on  evolutionary  lineages. 
Merychyus  cf.  M.  minimus  from  Boron  is  assignable  to  the 
genus  Merychyus  on  the  basis  of  the  large  orbit,  presence  and 
size  of  facial  fossa  and  prelacrimal  vacuity,  relatively  high- 
crowned  teeth  with  large,  posteriorly  inclined  upper  pre- 
molars, and  small  upper  C and  P,  (P,  is  the  lower  caniniform 
tooth  in  oreodonts). 

Based  on  length  of  upper  dentition,  the  Boron  species  is 
1 5 percent  smaller  than  M.  ( Metoreodon ) relictus  but  only  3 
percent  smaller  than  M.  ( Metoreodon ) relictus  fletcheri  Schultz 
and  Falkenbach,  1947,  from  the  Barstow  Formation.  The 
subgenus  Metoreodon,  based  on  M.  ( Metoreodon ) relictus,  is 
distinguished  from  the  subgenus  M.  ( Merychyus ) primarily 
by  its  more  complex  and  crowded  premolars,  P2^,  well  grooved 
externally,  P,_3  set  obliquely  in  the  jaw,  teeth  higher  crowned, 
and  several  other  skull  characters  not  preserved  in  the  Boron 
sample.  Merychyus  cf.  M.  minimus  from  Boron  has  a shallow 
lacrimal  fossa  and  fairly  high  crowned  teeth,  but  it  lacks  the 
premolar  complexity  and  crowding  typical  of  M.  ( Metoreo- 
don). 

The  length  of  the  upper  dentition  of  the  Boron  Merychyus 
is  25  percent  smaller  than  Ticholeptus  calimontanus  (Dough- 
erty, 1940),  from  the  Temblor  Formation,  but  it  is  as  high 
crowned.  The  genus  Ticholeptus  Cope,  1878,  although  placed 
in  a different  subfamily,  is  very  similar  to  Merychyus,  but 
differs  by  having  a proportionally  smaller  orbit,  a deeper 
jugal,  larger  upper  canine  and  P,,  and  more  anterior  infraor- 
bital foramen  above  P3-4. 

Merychyus  cf.  M.  minimus  from  Boron  is  larger  and  more 
hypsodont  than  M.  calaminthus  (type  series)  and  material 
referred  to  M.  calaminthus  from  the  type  area  of  the  Hector 
Formation  (specimen  UCR  10914  — Woodbume  et  al.,  1974), 
a specimen  from  the  Hector  Formation  in  the  northern  Cady 
Mountains  (specimen  UCR  10840— Miller,  1980)  and  a 
specimen  from  the  Orocopia  Mountains  (specimen  LACM 
27026  — Woodbume  and  Whistler,  1973).  In  overall  size,  the 
Boron  material  falls  between  M.  calaminthus/M.  crabilli  and 
M.  minimus  but  it  is  closer  to  the  latter.  Stevens  (in  Wood- 
bume etal.,  1974:19)  has  suggested  that  M.  calaminthus  and 
M.  crabilli  are  the  same  species,  a suggestion  she  reaffirmed 
in  a later  work  (Stevens,  1977:37),  but  stopped  short  of  pro- 
posing a formal  synonymy,  thus  they  are  considered  distinct 
in  this  study. 

Stevens  (1977)  has  suggested  two  evolutionary  trends  in 
Merychyus— decrease  in  “premolar  diameter”  (the  ratio  of 
the  length  of  the  Pi_4  to  the  length  of  M1-3)  and  an  increase 
in  hypsodonty.  The  material  from  Boron  (especially  UCR 
10319)  has  large  premolars  that  are  less  reduced  than  in  M. 
calaminthus  or  M.  crabilli,  but  has  cheek  teeth  that  are  more 
hypsodont  than  even  higher  crowned  individuals  of  M.  min- 
imus and  Merychyus  elegans  Leidy,  1858.  The  jugal  depth 
(as  a ratio  of  tooth  row  length)  is  shallow,  more  similar  to 
that  of  M.  calaminthus  and  M.  crabilli  than  that  of  M.  min- 
imus or  M.  elegans. 

Merychyus  cf.  M.  minimus  from  Boron  appears  distinct 
from  M.  calaminthus  and  M.  crabilli,  but  it  falls  within  the 


Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna  21 


22  Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna 


Figures  46-49.  IMerychyus  sp.  Figures  46-47,  UCR  10331,  adult  palate  with  well  worn  P2-M3  and  part  of  right  side  of  face;  (46)  right  lateral  view,  (47)  occlusal  view.  Figures 
48-49,  UCR  10332,  lower  mandible  with  alveolus  of  I ,_3,  partially  complete  C,  P,_3,  complete  P4-M3;  (48)  right  lateral  view,  (49)  occlusal  view.  All  actual  size. 


Table  4.  Selected  measurements  of  cf.  Hesperocamelus  sp.  and  an  indeterminate  species  of  small  camel  from  the  Boron  Local  Fauna.  Abbre- 
viations: AP  = anteroposterior  dimension  of  tooth,  TR  = width  of  tooth  at  anterior  loph(id),  * = approximate. 


cf.  Hesperocamelus 

Camelidae,  small  sp. 

UCR 

10366 

UCR 

10367 

UCR 

10365 

Width  palate,  anterior  end  P2 

27.3 

Length  P2-M3 

115* 

Length  dP2-M,  47.0 

P2  AP 

9.0 

M,  AP  16.2 

TR 

L6.1,  R6.3 

TR  6.2 

P3  AP 

L16.0,  R16.5 

Length  dP2  5.9 

TR 

9.8 

Length  dP,  9.0 

M1  AP 

19.6 

Length  dP4  16.8 

TR 

16.3 

M2  AP 

L27.7,  R27.7 

TR 

L21.1,  R22.0 

M3  AP 

29.3* 

TR 

23.5* 

M3  AP 

29.9 

TR 

10.0 

Width  distal  radioulna 

50.9  (UCR  10384) 

35.0  (UCR  10374) 

47.7  (UCR  10385) 

Width  proximal  metacarpal 

31.3  (UCR  21189) 

Width  distal  tibia 

42.0  (UCR  10339) 

34.0  (UCR  10371) 

42.0  (UCR  10387) 

37.0  (UCR  10388) 

Width  proximal  metatarsus 

27.9  (UCR  10376) 

Length  calcaneum 

1 17.7  (UCR  21186) 

69*  (UCR  10379) 

Length  astragalus 

48.0  (UCR  10392) 

36.8  (UCR  10378) 

considerable  range  of  variation  of  M.  minimus  in  many  char- 
acters. Merychyus  cf.  M.  minimus  from  Boron  is  probably 
distinct  from  M.  minimus,  but  larger  samples  and  more  com- 
pletely preserved  material  are  needed  before  this  can  be  con- 
clusively demonstrated. 

? Merychyus  sp. 

Figures  46-49,  Table  3 

MATERIAL.  Adult  palate  with  well-worn  P2-M3  and  part 
of  right  side  of  face,  UCR  10331;  lower  mandible  bearing 
alveolus  of  I,_3,  partially  complete  C and  P,_3,  complete  P4- 
M3,  UCR  10332;  fragment  of  left  dentary  with  P4-M,,  UCR 
10333,  fragment  of  right  dentary  with  M3,  UCR  21270. 

DESCRIPTION.  Size  similar  to  Merychyus  cf.  M.  mini- 
mus described  above;  jugal  deep;  infraorbital  foramen  above 
posterior  root  of  M1,  suggesting  a shorter  face;  upper  molars 
broad;  M3  with  split  metastyle  and  prominent  posterior  cin- 
gulum on  metaconule;  ratio  of  length  of  P‘~4  to  length  of 
M'~3  approximately  0.73,  considerably  smaller  than  the 
Merychyus  cf.  M.  minimus  described  above;  lower  premolars 
crowded,  P3  10  percent  larger  and  more  complex  than  Mer- 
ychyus cf.  M.  minimus  described  above. 


DISCUSSION.  Although  these  three  specimens  may  sim- 
ply be  larger,  shorter  faced,  more  robust  individuals  of  Mer- 
ychyus cf.  M.  minimus  described  above,  they  are  separated 
because  of  the  more  robust  teeth,  deep  jugal,  and  large  P3. 
Specimen  UCR  10322  is  a young  adult  which  may  account 
for  the  crowding  of  the  lower  premolars.  The  infraorbital 
foramen  is  clearly  more  posteriorly  located  than  in  the  three 
specimens  of  Merychyus  cf.  M.  minimus  described 
above,  suggestive  of  the  condition  found  in  species  of  Bra- 
chycrus  Matthew,  1901.  None  of  the  Boron  specimens  pre- 
serves the  nasal  region  of  the  cranium  which  is  diagnostic  of 
Brachycrus.  The  split  metastyle  and  strong  posterior  cingu- 
lum of  M3  and  jugal  depth  are  variable  characters  in  ore- 
odonts  (Lander,  personal  communication).  The  jugal,  al- 
though deep  for  species  of  Merychyus,  is  nowhere  near  as 
deep  as  in  Brachycrus,  but  is  similar  to  species  referred  to 
Ticholeptus.  The  Boron  specimens  are  much  smaller  than 
any  species  of  Brachycrus,  but  only  slightly  smaller  than  some 
specimens  referred  to  Ticholeptus  calimontanus. 

These  three  specimens  may  be  variants  of  the  associated 
Merychyus  cf.  M.  minimus,  but  there  is  more  difference  be- 
tween them  and  the  Merychyus  cf.  M.  minimus  than  between 
the  latter  and  M.  minimus  from  Nebraska.  Thus,  they  are 


Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna  23 


considered  distinct  until  larger  samples  are  available  to  more 
clearly  demonstrate  individual  variation.  There  is  some  sug- 
gestion of  convergence  with  features  characteristic  of  both 
Brachycrus  and  Ticholeptus. 

Family  Camelidae 
Subfamily  Aepycamelinae 
Hesperocamelus  Macdonald,  1 949 

cf.  Hesperocamelus  sp. 

Table  4 

MATERIAL.  Palate  with  RP‘~2,  RM2  and  part  of  RM!, 
LP2~3,  LM1"3,  isolated  ? upper  incisor,  UCR  10366;  LM3, 
UCR  10367;  posterior  portion,  RM„  UCR  10368;  associated 
left  distal  radioulna,  scaphoid  and  lunar,  UCR  10384;  distal 
epiphysis,  left  radioulna,  UCR  10385;  right  distal  tibia,  UCR 
10388;  two  left  distal  tibiae,  UCR  10387  and  UCR  10389; 
left  astragalus,  UCR  1 0392;  left  calcaneum,  UCR  2 1 1 86;  left 
proximal  metacarpal,  UCR  21 187;  right  proximal  metatar- 
sal, UCR  21  188;  proximal  phalanx,  UCR  10391. 

DESCRIPTION.  Size  (length  of  dentition  from  P1  to  M3) 
larger  than  the  aepycamelines  Oxydactylus  Peterson,  1904, 
and  Paratylopus  Matthew,  1904,  the  camelines  Dyseotylo- 
pus  Stock,  1935,  and  Priscocamelus  Stevens,  1969,  and  most 
protolabines;  about  same  size  as  the  aepycameline  Hespero- 
camelus alexandrae  (Davidson,  1923);  smaller  than  smallest 
species  of  Aepycamelus  Macdonald,  1956;  rostrum  fairly  nar- 
row at  anterior  root  of  P2  but  not  constricted  as  in  protola- 
bines; teeth  relatively  low  crowned  with  80-90  percent  of 
external  enamel  of  paracone  of  M3  exposed  when  this  tooth 
enters  occlusion;  premolars  large  in  comparison  to  molars; 
P1  large,  blade-like,  two  rooted;  P2  long,  blade-like,  with 
weak,  discontinuous  lingual  cingulum;  P3  long,  parastyle  large, 
lingual  cingulum  well  developed,  with  distinct  central  gap; 
M'~3  with  weak,  but  distinct  vertical  ribs  on  paracone,  ribs 
nearly  lacking  on  metacone,  metastyle  and  parastyle  strong; 
M3  not  elongated  anteroposteriorly  compared  to  M1;  M3  with 
strong  metastylid  and  reduced  hypoconulid;  metacarpals  and 
metatarsals  fused;  calcaneum  long  and  slender. 

DISCUSSION.  Camel  taxonomy  is  usually  based  on  as- 
sociated skulls  and  postcranial  material.  Tentative  associa- 
tion of  the  palate  UCR  20366  with  some  of  the  postcranial 
material  provides  for  a composite  picture  of  this  medium 
sized  camel.  The  palate  is  broken  anterior  to  the  P2  so  that 
the  degree  of  palatal  constriction  and  length  of  the  rostrum 
are  lacking  (key  characters  in  distinguishing  protolabines  from 
aepycamelines).  Although  the  palate  is  fairly  narrow  between 
the  anterior  roots  of  the  P2,  these  teeth  are  not  turned  inward 
anteriorly  as  is  typical  in  protolabines. 

The  premolars  are  unreduced  and  large  in  comparison  to 
the  molars,  a feature  typical  of  many  Early  Miocene  camels, 
but  they  lack  the  robustness  typical  of  Miolabis  Hay,  1899 
(Aepycamelinae).  The  large,  blade-like  P'  is  similar  to  that 
of  most  protolabines  and  aepycamelines,  but  P1  in  Hespero- 
camelus is  typically  more  caniniform.  The  P2  in  the  cf.  Hes- 


perocamelus sp.  from  Boron  is  larger  in  comparison  to  the 
other  cheek  teeth  than  in  most  protolabines  or  aepycame- 
lines. The  P3  is  also  large.  The  discontinuous  lingual  cingulum 
of  the  P3  is  a variable  character  but  is  similar  to  the  primitive 
protolabine  Michenia  Frick  and  Taylor,  1971,  and  Hesper- 
ocamelus. The  crown  of  P4  is  missing  on  both  sides  of  the 
palate. 

The  molars  are  low  crowned  as  is  characteristic  of  aepy- 
camelines and  most  protolabines  and  in  contrast  to  the  higher 
crowned  cameline  Procamelus  Leidy,  1858,  and  the  notably 
high-crowned  stenomylines  (Frick  and  Taylor,  1968).  The 
vertical  ribs  on  the  paracone  and  metacone  are  more  reduced 
than  those  of  the  primitive  protolabines  Michenia  and  Tany- 
mykter  Honey  and  Taylor,  1978,  and  the  primitive  aepy- 
camelines Oxydactylus  and  Miolabis.  They  are  more  like 
those  of  stenomylines  and  the  more  derived  aepycamelines 
Hesperocamelus  and  Aepycamelus.  However,  the  strong  me- 
sostyle  is  more  like  that  of  Michenia,  Tanymykter,  and  Oxy- 
dactylus. The  M3  is  not  anteroposteriorly  lengthened  in  re- 
lation to  the  length  of  the  M1  as  is  common  in  many  derived 
protolabines.  Two  isolated  M,’s  bear  relatively  strong  meta- 
stylids  which  are  typical  of  most  protolabines  and  aepy- 
camelines, otherwise  these  referred  specimens  provide  little 
additional  information. 

Fragments  of  a fused  metacarpal  and  fused  metatarsal  oc- 
cur in  the  fauna,  but  neither  are  complete  enough  to  obtain 
an  estimate  of  their  length.  The  degree  of  fusion  distinguishes 
this  Boron  material  from  Miolabis,  Paratylopus,  most  species 
of  Oxydactylus  and  all  species  of  Michenia  except  M.  exilis 
(Matthew,  1 960)  all  of  which  have  unfused  metapodials.  There 
is  one  partial  calcaneum  which  is  quite  long  and  slender,  a 
condition  common  in  stilt-legged  (aepycameline)  camels. 

In  most  observable  characters,  cf.  Hesperocamelus  sp.  from 
Boron  is  not  clearly  distinguishable  from  early  protolabines 
( Michenia , Tanymykter)  or  early  aepycamelines  ( Oxydac- 
tylus).  As  clearly  pointed  out  by  Honey  and  Taylor  (1978: 
377),  it  is  difficult  to  distinguish  these  taxa  on  cheek  teeth 
alone.  However,  the  Boron  cf.  Hesperocamelus  sp.  seems  to 
lack  the  restricted  rostrum  characteristic  of  all  protolabines 
and  it  is  larger  than  any  species  of  Oxydactylus.  Thus,  it  is 
referred  to  the  larger  aepycameline  with  fused  metapodials, 
Hesperocamelus.  Except  for  the  blade-like  P1  and  slightly 
smaller  size,  it  closely  resembles  Hesperocamelus  alexandrae 
from  the  Barstow  Formation. 

Subfamily  undetermined 

Camelidae,  small  species 

Table  4 

MATERIAL.  Juvenile  dentaries  with  LdP2_4,  RdP4,  RM,, 
and  LM,,  UCR  10365;  proximal  right  metacarpals  III  and 
IV,  UCR  21189;  proximal  right  metatarsal,  UCR  10376; 
proximal  right  scapula,  UCR  10370;  proximal  end  right  ra- 
dioulna, UCR  10371;  proximal  end  left  radioulna,  UCR 
10372;  proximal  articular  facet  right  radioulna,  UCR  10373; 
distal  epiphysis  right  radioulna,  UCR  10374;  right  lunar  UCR 
21190;  right  entocuneiform,  UCR  10382;  distal  left  tibia, 
UCR  10377;  right  malleolus,  UCR  21191;  right  juvenile  cal- 


24  Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna 


caneum,  UCR  10379;  right  astragalus,  UCR  10378;  two  right 
cuboids,  UCR  10393  and  UCR  10394;  right  navicular,  UCR 
10395;  proximal  end,  proximal  phalanx,  UCR  10381. 

DISCUSSION.  This  small  camel  is  represented  by  a pair 
ofjuvenile  dentaries  with  the  M,  the  only  erupted  permanent 
tooth  and  with  the  dP3_4  moderately  worn.  The  specimen  is 
broken  at  the  symphysis,  but  there  is  an  alveolus  for  a large 
P,.  Additional  small,  postcranial  camelid  elements  may  also 
belong  to  this  taxon.  One  fragment  represents  the  proximal 
ends  of  metacarpals  III  and  IV  which  are  closely  appressed 
but  unfused;  a fragment  of  the  proximal  end  of  the  metatarsal 
is  fused. 

The  material  is  too  incomplete  to  permit  even  a subfamilial 
assignment,  but  clearly  demonstrates  the  presence  of  a small 
camel  in  the  fauna.  Based  on  postcranial  elements,  this  is  the 
most  common  camel.  This  small  camel  could  be  any  of  sev- 
eral protolabines  that  are  common  in  Miocene  assemblages 
of  the  western  United  States  or  a species  of  Oxydactylus. 
Priscocamelus  or  Paratylopus,  all  less  common. 

Camelidae,  large  species 

DISCUSSION.  A large  camel  is  represented  by  a left  distal 
femur  fragment,  UCR  10386,  and  a left  proximal  scapular 
fragment,  UCR  10383.  These  specimens  suggest  an  animal 
about  half  again  as  large  as  the  species  referred  to  cf.  Hes- 
perocamelus  sp.  They  could  represent  one  of  several  small 
species  of  Aepycamelus  which  are  common  in  Miocene  as- 
semblages of  the  Great  Basin. 

Family  Palaeomerycidae 

Subfamily  Aletomerycinae 
Aletomeryx  Lull,  1920 

Aletomeryx  occidentalis  new  species 

Figures  51-59,  Table  5 

HOLOTYPE.  UCR  10335,  partial  young  adult  right  den- 
tary with  P2  (natural  cast  in  part)  and  complete  P3  through 
M3. 

TYPE  LOCALITY.  UCR  locality  RV  6421  (see  complete 
description  under  Cupidinimus  boronensis  above). 

REFERRED  MATERIAL.  Portion  of  right  frontal  and 
horn  base,  UCR  10348;  partial  left  dentary  with  I2,  well  worn 
P2-M2  and  roots  for  I,,  I3,  and  C,  UCR  10336;  partial  right 
dentary  with  little  worn  P2-M2,  UCR  10337;  partial  sub- 
adult right  dentary  with  dP2,  unworn  P3-M2,  UCR  10338; 
fragment  of  right  dentary  with  moderately  worn  M2_3,  UCR 
10339;  fragment  of  right  dentary  with  moderately  worn  M„ 
UCR  10340;  partial  right  juvenile  dentary  with  symphysis, 
early  wear  dP2_4,  M,,  UCR  10341;  partial  right  juvenile  den- 
tary with  early  wear  dP2_.„  M,  UCR  10342;  partial  left  ju- 
venile dentary  with  early  wear  dP2_^,  UCR  10343;  partial 
right  maxilla  with  early  wear  M 1-3,  UCR  10344;  partial  young 
adult  right  maxilla  with  alveolus  for  P1-2,  partially  complete 
P4  and  M1-2;  UCR  10345;  isolated  fragment  LdP2,  UCR 
10347;  isolated  fragment  ?RP2,  UCR  10398;  left  proximal 
scapula,  UCR  10352;  two  right  distal  humeri,  UCR  10356 


mesostyle 
metacone 
metastYle^^. 


metaconule 


paracone  metacomd  parastylid 


entostyle 


entoconid/  / 
entostylid 


metaconid 


I / paraconid 
/ / ...parastylid 


protoconid 


hypocomd 


Figure  50.  Tooth  cusp  terminology  used  in  Aletomeryx. 


and  UCR  10357;  left  proximal  radius,  UCR  10351;  distal 
half  right  radioulna,  UCR  10349;  associated  distal  end  left 
radioulna,  scaphoid,  lunar,  unciform,  fused  magnum-trap- 
ezoid, proximal  metacarpal,  UCR  1 0350;  left  cuneiform,  UCR 
21192;  associated  right  tibia  lacking  proximal  articulation, 
astragalus,  calcaneum,  naviculocuboid,  entocuneiform,  and 
proximal  fused  metatarsals  III  and  IV,  UCR  21 193;  four  left 
distal  tibiae,  UCR  10358,  UCR  10359,  UCR  10360,  and 
UCR  10364;  right  calcaneum,  UCR  10361;  associated  left 
naviculocuboid,  entocuneiform,  fused  metatarsals  III  and  IV, 
vestiges  of  metatarsus  II  and  V,  UCR  10355;  two  left  na- 
viculocuboids,  UCR  10362  and  UCR  10363;  left  entocu- 
neiform, UCR  10354;  proximal  phalanx  UCR  21 194. 

DIAGNOSIS.  Slightly  larger  (1.5  percent)  than  Aletome- 
ryx marslandesis  Frick,  1937,  the  largest  species  of  Aleto- 
meryx, but  smaller  (about  2 percent  than  the  smallest  species 
of  Sinclairomeryx  Frick,  1937;  horn  Aletomeryx- like,  situ- 
ated over  posterior  part  of  orbit  with  slight  forward  tilt;  den- 
tition hypsobrachyodont;  P2  large  for  species  of  Aletomeryx; 
premolars  with  simple  pattern  compared  to  common  pa- 
laeomerycids  such  as  Dromomeryx  Douglass,  1 909,  but  more 
complicated  pattern  than  that  of  described  species  of  Ale- 
tomeryx; lower  premolars  with  relatively  straight  crests  and 
lacking  expanded  metaconid;  posterior  labial  comer  (hypo- 
stylid)  of  P3_4  expanded  into  distinct  cuspid  that  connects  to 
hypoconid  by  short  crest  in  moderate  wear;  labial  end  of 
central  crest  (metaconid)  slightly  expanded  anterolabially  in 
P4  but  much  less  developed  than  in  dromomerycines  and 
giraffids;  opening  of  lingual  flexids  of  P4  not  directed  poste- 
riorly as  in  dromomerycines;  M,_,  with  well-developed  an- 
terior cingula  and  anterolabial  buttresses  which  join  in  wear; 
hypoconulid  of  M3  formed  by  closed  lingual  crescent  or  cres- 
cent in  combination  with  smaller  lingual  cusp;  post-sym- 
physial  diastema  short;  limbs  lightly  constructed,  proximal 
ends  of  metatarsals  II  and  V retained  as  vestiges. 

ETYMOLOGY.  From  occidentalis  Latin,  of  the  west,  in 
reference  to  the  occurrence  of  this  species  in  far  western  range 
of  distribution  of  the  genus. 

DESCRIPTION.  Orbital  Horn.  Single  broken  pedicel, 
UCR  10348,  situated  over  posterior  portion  of  orbit,  with 
slight  forward  tilt  and  some  suggestion  of  forward  curves; 
external  central  ridge,  leading  to  postorbital  process  thinner 
than  in  the  small  species,  Aletomeryx  gracilis  Lull,  1920; 
cross  section  of  hom  pedicel  at  base  broadly  triangular  in 
outline;  internal  surface  broadly  convex,  not  flat  as  Sinclai- 


Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna  25 


Table  5.  Selected  measurements  of  Aletomeryx  occidentalis  n.  sp.  Abbreviations:  AP  = anteroposterior  dimension  of  tooth,  TR  = width  of 
tooth  at  anterior  loph(id),  CH  = crown  height  at  paracone  in  upper  molars,  at  protoconid  in  lower  premolars,  at  metaconid  in  lower  molars 
(figures  given  only  for  unworn  specimens),  * = approximate. 


UCR  UCR  UCR  UCR  UCR  UCR  UCR  UCR 

Dentary  10335  10336  10337  10338  10339  10341  10342  10343 


Length  P2-M, 

75.2 

69.0 

Length  P-P, 

24.0 

23.9 

27.4 

25.0 

Length  M,-M3 

48.8 

45.7 

P2 

AP 

5.5* 

6.4 

7.8 

7.0 

TR 

3.0* 

3.5 

3.7 

3.5 

P3 

AP 

8.4 

7.5 

9.1 

8.0 

TR 

4.8 

4.4 

4.9 

4.7 

CH 

6.7* 

7.7 

P< 

AP 

9.9 

10.0 

1 1.1 

10.1 

TR 

6.0 

5.2 

5.0* 

4.5 

CH 

10.2 

10.5 

M, 

AP 

1 1.5 

9.1 

14.0 

13.8 

13.1 

TR 

7.0 

6.2 

6.5 

6.5 

6.2 

CH 

8.6 

m2 

AP 

14.2 

14.0 

16.0 

17* 

15* 

TR 

8.8 

8.0 

8.5 

8.0 

- 

CH 

13.1 

1 1.7 

M, 

AP 

20.0 

21.7 

21.0 

TR 

8.8 

8.1 

8.4 

CH 

12.2 

Length  dP,-dP4 

28.0 

26.6 

27.0 

dP2  AP 

6* 

5.5 

5.4 

TR 

3.1 

2.6' 

dP,  AP 

7.0 

6.1 

8.0 

TR 

3.6 

4.8 

4.0 

dP4  AP 

14.4 

14.1 

12.5 

TR 

5.4 

5.8 

5* 

UCR 

UCR 

UCR 

Maxilla 

10344 

10345 

10347 

Length  P2-M2 

# 

00 

IT) 

Length  P2-P4 
P2  AP 
TR 

28* 

P4  AP 

9.1 

TR 

9.5' 

M1  AP 

12* 

14.1 

TR 

11.2 

M2  AP 

14.6 

18.0 

TR 

14.0 

14.2 

CH 

15.0 

M3  AP 

14.8 

TR 

14.6 

CH 

12.0 

26  Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna 


Table  5.  Continued. 


Axial  skeleton 

UCR 

10349 

UCR 

10350 

UCR 

10356 

UCR 

10357 

Width  distal  humerus 

25.0 

23.3 

Width  distal  radioulna 

22.8 

21.4 

Width  proximal  metacarpal 

18.3 

UCR 

UCR 

UCR 

UCR 

UCR 

UCR 

UCR 

UCR 

10355 

10358 

10360 

10361 

10362 

10363 

10364 

21193 

Width  distal  tibia 

23.8 

23.5 

23.0 

21* 

Width  naviculocuboid 

20.0 

19.0 

21.4 

20.6 

Length  calcaneum 

54.6 

51* 

Length  astragalus 

23.8 

Width  prox.  metatarsal 

20.0 

19.0 

romeryx;  horn  pedicel  becomes  triangular  along  posterior 
border  above  base;  two  foramina  separated  by  a small  bridge 
of  bone  present  in  frontal  above  orbit;  suggestion  of  third 
foramen  at  frontal-lacrimal  suture. 

Upper  Dentition.  Enamel  more  rugose  than  in  lower  den- 
tition; styles  well  developed,  but  not  greatly  enlarged;  alveo- 
lus for  P2  in  UCR  10345  suggests  broader  tooth  than  in  other 
species  of  Aletomeryx;  alveolus  for  P3  about  same  size  as  in 
other  species  of  Aletomeryx  or  Sinclairomeryx;  P2  in  UCR 
10398  with  well  developed  lingual  cingulum;  P4  only  partially 
preserved,  but  with  fossette  divided  posteriorly;  M1-3  with 
well-developed  styles  and  accessory  folds  in  fossettes;  weak 
anterior  cingulum;  small  pillar  (entostyle)  in  M1-2  lacking  in 
M3. 

Upper  Deciduous  Dentition.  dP2  similar  to  permanent  P2 
with  single  high  labial  crest;  continuous  lingual  cingulum 
around  anterior  end  of  tooth  with  shallow  groove  below  high 
portion  of  labial  crest  (protocone);  small  spur  leading  from 
center  of  anterior  portion  of  cingulum  to  base  of  protocone. 

Dentaries  and  Lower  Dentition.  Lower  jaws  lightly  built, 
with  slight  inward  curve  in  region  of  symphysis;  post-sym- 
physial  diastema  proportionally  shorter  than  in  smaller  species 
of  Aletomeryx ; molars  high  crowned  with  less  than  half  the 
entire  crown  height  of  M3  exposed  when  this  tooth  comes 
into  occlusion;  enamel  lightly  rugose;  premolars,  particularly 
P2,  less  reduced  than  in  other  described  species  of  Aletomeryx 
or  Sinclairomeryx. 

I , 3 and  lower  canine  all  about  the  same  size  judging  from 

roots,  all  directed  anteriorly. 

P2  (based  on  UCR  10327  and  UCR  10336)  larger,  with 
more  complicated  pattern  than  in  previously  known  Aleto- 
meryx species;  with  well-developed  anterior  cusp  (paraconid) 
that  persists  in  wear;  paraconid  connected  by  labial  crest  to 
high  central  cusp  (protoconid);  separate  entoconid  and  en- 
tostylid  present  in  early  wear. 

P3  with  more  complicated  cusp  pattern  than  P2;  paraconid 
and  parastylid  well  developed,  becoming  more  separate  near 
base  of  crown;  protoconid  most  prominent  cusp;  entoconid 
and  entostylid  well  developed  with  intervening  flexid  opening 


more  labially  than  in  most  Aletomeryx  species;  suggestion  of 
low  postero-labial  enlargement  (hypostylid)  which  occludes 
only  in  late  stage  of  overall  tooth  wear. 

P4  with  separate  and  distinct  paraconid  and  parastylid; 
parastylid  an  isolated  cusp  in  early  wear;  metaconid  begin- 
ning as  isolated  cusp  but  strongly  connected  to  protoconid 
by  central  crest;  metastylid  absent;  posterior  portion  of  P4 
distinctly  bifurcate  with  separate  entoconid  and  entostylid; 
flexid  between  entoconid  and  entostylid  opens  more  poste- 
riorly than  in  P3;  posterolabial  comer  (hypostylid)  well  de- 
veloped to  mid  height  of  tooth  crown. 

M,  small  in  proportion  to  P4  compared  to  other  species  of 
Aletomeryx  or  Sinclairomeryx;  parastylid,  anterior  cingu- 
lum, and  anterolabial  buttress  (protostylid)  well  developed. 

M2  with  well-developed  anterior  cingulum  bearing  distinct 
anterolabial  cusp  or  buttress  (protostylid)  in  unworn  tooth, 
cusp  joins  cingulum  after  moderate  wear,  persists  to  late 
wear;  suggestion  of  metastylid  present  as  inflection  in  enamel 
in  UCR  10337  and  UCR  10338,  but  not  persistent  in  wear; 
ectostylid  developed  as  small,  low  pillar;  connection  between 
posterior  end  of  entoconid  and  posterolingual  extension  of 
hypoconid  (=  entostylid)  delayed  until  moderate  wear,  form- 
ing a shallow  enamel  infolding  in  posterolingual  comer  of 
tooth. 

M3  with  hypoconulid  formed  by  completely  closed  labial 
loop  in  the  type,  UCR  10335,  and  UCR  10336;  UCR  10339 
(Figures  47-48)  with  additional  small,  lingually  projected 
spur  and  small  lingual  cusp;  anterior  cingulum  not  as  well 
developed  as  in  M,  or  M2;  ectostylids  variably  developed 
between  protoconid  and  hypoconid,  hypoconid  and  hypo- 
conulid. 

Lower  Deciduous  Dentition.  dP2  very  similar  to  the  P2, 
except  with  deeper  posterior  flexid  more  persistent  with  wear; 
dP3  with  better  developed  paraconid,  parastylid,  and  bifur- 
cation of  the  anterior  cingulum;  posterior  flexid  of  dP3  open- 
ing more  directly  lingually  than  in  P3,  with  small  cusp  in 
anterior  labial  comer  (protostylid)  that  forms  anteriorly 
opening  flexid  after  moderate  wear;  dP4  three  lobed,  resem- 
bling reversed  M3,  but  with  stronger  ectostylids. 


Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna  27 


Figures  51-59.  Aletomeryx  occidentalis  n.  sp.  Figures  51-52,  Holotype,  UCR  10335,  right  adult  mandible  with  early  wear  P2  (in  part  natural 
cast)  and  complete  P3-M3;  (51)  labial  view,  (52)  occlusal  view.  Figures  53-54,  UCR  10339,  RM3;  (53)  lingual  view,  (54)  occlusal  view.  Figures 
55-56,  UCR  10342,  juvenile  right  dentary  fragment  with  early  wear  RdP2_4,  part  of  M,;  (55)  labial  view,  (56)  occlusal  view.  Figure  57,  UCR 
10348,  right  horn  fragment.  Figures  58-59,  UCR  10344,  RM2-3;  (58)  labial  view,  (59)  occlusal  view.  All  actual  size. 


Limbs.  Long  and  lightly  built;  front  and  hind  metapodials 
well  fused;  principal  metatarsal  with  vestiges  of  proximal 
ends  of  both  metatarsals  II  and  V;  vestigial  metatarsal  II 
elongate,  triangular,  fused  to  central  metatarsal;  vestigial 
metatarsal  V stubby,  broad,  unfused;  vestige  of  metacarpal 


II  also  present  as  suggested  by  flattened,  rugose  area  on  meta- 
carpal III. 

Discussion.  The  dentition  of  Aletomeryx  occidentalis  is 
high  crowned,  more  so  than  most  taxa  referred  to  Palaeo- 
merycidae.  The  high-crowned  teeth  and  narrow  P,_4  distin- 


28  Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna 


guish  Aletomeryx  occidentalis  from  all  the  more  deer-  or 
giraffe-like  palaeomerycids:  Dromomeryx,  Rakomeryx  Frick, 
1937,  Cranioceras  Matthew,  1918,  Barbouromeryx  Frick, 
1937,  Drepanomeryx  Sinclair,  1915,  and  Yumaceras  Frick, 
1937,  of  North  America,  Palaeotneryx  Von  Meyer,  1834, 
Procervulus  Gaudry,  1878,  Heterocemas  Young,  1937,  and 
Triceromeryx  Villalta,  Crusafont  and  Lavocat,  1946,  from 
Europe  and  Asia  and  Climacoceras  Maclnnes,  1936,  Can- 
thumeryx  Hamilton,  1973,  and  Propataeoryx  Slromer,  1926, 
from  Africa.  Aletomeryx  occidentalis  is  distinguished  from 
the  smaller,  hornless,  North  American  leptomerycine  Lep- 
tomeryx  Leidy,  1853,  Pseudoparablastomeryx  Frick,  1937, 
and  Pronodens  Koemer,  1940  (see  Taylor  and  Webb,  1976), 
from  the  blastomerycines  Blastomeryx Cope,  1877,  and  Ma- 
chaeromeryx  Matthew,  1926,  and  from  the  palaeomerycid 
Longirostromeryx  Frick,  1937,  by  the  high-crowned  teeth 
and  presence  of  orbital  horns. 

Only  three  palaeomerycids,  Aletomeryx,  “ Dyseomeryx,” 
and  Sinclairomeryx,  have  high-crowned,  antilocaprid-like 
teeth.  The  relatively  large  premolars,  particularly  P2,  and  the 
greater  complexity  of  P3_4,  with  a well-developed  metaconid 
on  P4  and  hypostylid  on  P3_4,  are  characters  derived  with 
respect  to  any  previously  described  species  of  Aletomeryx  or 
Sinclairomeryx. 

There  is  little  clear  distinction  between  species  of  Aleto- 
meryx, “ Dyseomeryx ,"  and  Sinclairomeryx  because  many 
characters  overlap.  However,  the  structure  of  the  horn  ped- 
icel, the  development  of  the  anterolabial  buttress  and  anterior 
cingulum  on  M2,  and  the  hypoconulid  crescent  of  M3  favor 
assignment  of  the  Boron  species  to  the  genus  Aletomeryx. 
Some  smaller  specimens  of  Aletomeryx  gracilis  approach  the 
crown  height  of  Aletomeryx  occidentalis,  but  these  lack  the 
lower  premolar  complexity. 

There  is  also  confusion  as  to  the  validity  of  the  generic  (or 
subgeneric)  names  Aletomeryx,  Dyseomeryx,  and  Sinclai- 
romeryx within  the  Subfamily  Aletomerycinae.  According  to 
Stirton  (1944)  in  his  sort  discussion  of  the  Family  Palaeo- 
merycidae,  the  genus  Aletomeryx  Lull,  1 920,  is  distinct  but 
the  genera  (or  subgenera)  Dyseomeryx  Matthew,  1924,  and 
Sinclairomeryx  Frick,  1937,  are  synonymous,  Dyseomeryx 
with  priority.  He  did  not  thoroughly  discuss  his  reasons  for 
this  synonymy,  however.  On  the  other  hand,  Frick  (1937) 
stated  that  Dyseomeryx  is  in  part  synonymous  with  Aleto- 
meryx. 

Matthew  (1924:196)  defined  the  subgenus  Dyseomeryx, 
choosing  Blastomeryx  marshi  Lull,  1920,  as  the  type,  which 
then  became  Blastomeryx  {Dyseomeryx)  marshi  (Lull)  as  re- 
combined by  Matthew  (1924).  But,  according  to  Frick  (1937: 
152),  Blastomeryx  {Dyseomeryx)  marshi  (Lull)  actually  be- 
longs in  the  genus  Aletomeryx,  and  therefore  Blastomeryx 
(Dyseomeryx)  marshi  became  Aletomeryx  marshi  (Lull)  as 
recombined  by  Frick  (1938).  Because  Frick  (1937)  found  it 
necessary  to  refer  the  type  of  the  subgenus  (Dyseomeryx)  to 
Aletomeryx,  while  he  considered  other  forms  referred  to  (Dy- 
seomeryx) generically  distinct,  he  defined  the  new  genus  Sin- 
clairomeryx, naming  as  the  type  species  Sinclairomeryx  sin- 
clairi  Frick,  1937.  Frick  also  questionably  assigned  another 
of  Matthew’s  species,  Blastomeryx  (Dyseomeryx)  riparius 


Matthew,  1924,  to  Sinclairomeryx.  The  type  of  this  species 
is  a left  maxilla  fragment,  and  certain  assignment  to  Sin- 
clairomeryx by  Frick  would  necessitate  an  associated  horn 
core,  which  was  not  present.  Based  on  comparisons  of  den- 
titions, later  workers  (Skinner  et  al.,  1977)  have  indicated  5. 
riparius  and  S.  sinclairi  are  synonymous,  with  S.  riparius 
having  priority. 

Although  I have  not  reviewed  the  original  material,  it  would 
seem  that  Frick  (1937)  substantiated  his  view  and  I have 
accepted  the  genera  Sinclairomeryx  (=  Dyseomeryx  in  part) 
and  Aletomeryx. 

AGE  AND  BIOSTRATIGRAPHY 
RADIOMETRIC  CORRELATION 

There  are  three  radiometric  dates  on  different  outcrops  of 
the  Saddleback  Basalt:  18.3  ± 0.6  ma,  19.7  ± 0.6  ma  (R.B. 
Kistler,  personal  communication),  and  20.3  ± 0.7  ma  (Arm- 
strong and  Higgens,  1973).  The  Saddleback  Basalt  occurs 
146  m below  the  Boron  Local  Fauna  in  the  open  pit  mine 
and  thus  these  radiometric  dates  provide  a maximum  age 
for  the  fossil  assemblage.  Based  on  biostratigraphic  corre- 
lations (see  below),  the  Boron  Local  Fauna  is  distinctly  older 
than  the  typical  fossil  vertebrate  assemblages  of  the  Barstow 
Formation  which  have  a maximum  age  of  at  least  16.3  ± 
0.3  ma  (Woodbume  and  Tedford,  1982)  and  the  Early  Bar- 
stovian  Dry  Canyon  fauna  of  the  Caliente  Formation  dated 
at  16.5  ± 1.3  ma  (Turner,  1970,  KA  2127).  These  dates 
provide  a minimum  age  for  the  Boron  Local  Fauna.  On 
biostratigraphic  grounds,  the  Boron  Local  Fauna  appears  close 
to  the  Phillips  Ranch  Local  Fauna  which  is  underlain  by  a 
17.6  ma  date  (Evemden  et  al.,  1964,  KA  478).  All  these  form 
a consistent  pattern  with  the  Boron  Local  Fauna  falling  be- 
tween approximately  16.5  and  18.5  ma. 

BIOSTRATIGRAPHIC  CORRELATIONS 

Generally,  correlation  of  southern  Californian  Arikareean 
and  Hemingfordian  faunas  with  the  classical  sequence  in 
Nebraska  and  neighboring  states  has  been  difficult  because 
the  fossil  assemblages  in  these  widely  separated  areas  usually 
have  had  little  in  common.  The  Boron  Local  Fauna  presents 
a rare  exception  to  this  generality  for  it  contains  two  taxa 
very  similar  to  species  in  Nebraska,  Merychyus  cf.  M.  min- 
imus and  Aletomeryx  occidentalis.  Merychyus  minimus,  re- 
stricted to  the  early  Hemingfordian  of  the  central  United 
States,  is  a quite  variable  species  and  M.  cf.  M.  minimus 
from  Boron  falls  within  this  variation.  A.  occidentalis,  al- 
though distinct  at  the  species  level  and  probably  representing 
a separate  lineage  from  the  Aletomeryx/Sinclairomeryx  lin- 
eage in  the  central  United  States,  is  close  to  A.  marslandensis 
from  the  early  Hemingfordian  of  Nebraska.  Taken  as  a 
whole,  the  Boron  Local  Fauna  is  best  considered  early 
Hemingfordian  in  age,  but  probably  near  the  transition  to 
the  late  Hemingfordian. 

Compared  to  mid-continent  assemblages,  the  Boron  Local 
Fauna  is  not  particularly  diverse,  and  may  lack  “key”  taxa 
which  were  not  preserved  or  may  not  have  been  present.  To 


Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna  29 


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30  Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna 


some  extent,  the  Boron  Local  Fauna  represents  a unique 
combination  of  taxa,  with  a rodent  ( Trogomys ) previously 
thought  to  be  Arikareean,  a palaeomerycid  genus  (Aleto- 
meryx)  and  an  oreodont  species  ( Merychyus  cf.  M.  minimus), 
elsewhere  restricted  to  the  early  Hemingfordian,  and  a di- 
podomyine  rodent  genus  (Cupidinimus)  and  camel  (cf.  Hes- 
perocamelus  sp.)  the  latter  genus  previously  restricted  to  the 
Barstovian  and  later. 

Absent  at  Boron  are  taxa  which  are  generally  typical  of 
late  Hemingfordian:  small  merychippines,  middle  sized  ore- 
odonts  larger  than  Merychyus  (Brachycrus,  Ticholeptus),  dro- 
momerycids  (Bouromeryx  Frick,  1937),  and  antilocaprids. 
Conclusions  based  on  absent  taxa  are  always  questionable, 
but  the  lack  of  these  taxa  found  in  common  association  in 
other  late  Hemingfordian  faunas  of  southern  California  and 
Nebraska  strongly  suggest  a real  absence  in  the  Boron  Local 
Fauna,  not  just  an  artifact  of  the  record  and  further  supports 
a slightly  older  age  than  late  Hemingfordian.  Camel  biostra- 
tigraphy is  not  well  documented,  and  thus  the  cf.  Hespero- 
camelus  sp.,  a genus  previously  restricted  to  Barstovian  and 
later,  is  not  considered  particularly  significant. 

The  microfauna,  although  seemingly  well  preserved,  lacks 
the  common  Arikareean/Hemingfordian  genus  Prohetero- 
mys  and  the  eomyid/geomyoid/zapodid  elements  common 
in  earlier  Miocene  assemblages.  Cupidinimus,  the  most  com- 
mon taxon  in  the  Boron  Local  Fauna  and  a common  rodent 
in  the  earliest  Barstovian,  is  absent  in  the  only  other  Hem- 
ingfordian microvertebrate  assemblage  in  southern  Califor- 
nia (Vedder  Local  Fauna;  Lindsay,  1 974)  and  thus  may  seem 
out  of  place  at  Boron.  However,  its  absence  may  be  envi- 
ronmentally controlled  and  its  presence  at  Boron  merely  pro- 
vides an  earlier  link  between  this  common  Barstovian  taxon 
and  its  presumed  ancestor,  Trogomys. 

The  Boron  Local  Fauna  displays  greatest  similarity  with 
the  central  United  States  fossil  assemblages  of  the  Running- 
water  Formation  (Cook,  1965;  McKenna,  1965),  Rosebud 
Formation  (Macdonald,  1963,  1970;  Macdonald  and  Hark- 
sen,  1968),  Batesland  Formation  (Harksen  and  Macdonald, 
1967;  Martin,  1976),  and  Martin  Canyon  “beds”  (Galbreath, 
1953;  Wilson,  1 960).  These  fossil  assemblages  are  considered 
typical  of  early  Hemingfordian  (Tedford  et  al.,  1973).  In 
addition  to  M.  minimus  and  Aletomeryx  species,  these  faunas 
also  contain  taxa  similar  to  the  Boron  forms  such  as  Ar- 
chaeolagus,  Hypolagus,  Miospermophilus,  Mookomys,  prim- 
itive aepycamelines  ( Oxydactylus ) and  primitive  protola- 
bines  ( Tanymykter , Protolabis),  even  though  none  of  these 
taxa  are  limited  to  the  early  Hemingfordian. 

There  is  less  similarity  between  the  Boron  Local  Fauna 
and  the  later  Hemingfordian  fossil  assemblages  of  the  Sheep 
Creek  Formation  (Skinner  et  al.,  1977)  and  the  Box  Butte 
Formation  (Galusha,  1975),  the  latter  containing  a diversity 
of  horses,  including  Merychippus  spp.,  larger  oreodonts  (Bra- 
chycrus, M.  ( Metoreodon ),  Ticholeptus),  Sinclairomeryx  spp. 
(but  no  Aletomeryx),  dromomerycines,  and  antilocaprids. 

The  Boron  Local  Fauna  can  also  be  correlated  with  south- 
ern California  fossil  assemblages  (Figure  60).  However,  such 
correlations  present  some  difficulty  because  of  the  seemingly 
unusual  assemblage  of  taxa  in  the  Boron  Local  Fauna.  The 


Arikareean/Hemingfordian  interval  is  not  well  represented 
in  southern  California.  Most  fossil  assemblages  referred  to 
this  time  span  are  poorly  preserved,  lack  taxonomic  diversity 
and  most  correlations  with  the  central  United  States  standard 
have  been  tenuous.  In  addition,  the  Boron  Local  Fauna  ap- 
pears to  sample  a restricted  time  interval  not  previously  seen 
in  southern  California  or  the  southern  Great  Basin. 

Most  Arikareean  fossil  assemblages  from  southern  Cali- 
fornia have  been  derived  from  sedimentary  basins  west  of 
the  San  Andreas  Fault,  which,  by  currently  accepted  plate 
tectonic  models,  places  these  basins  from  100-300  km  to  the 
southeast  at  their  time  of  development  (see  Crowell,  1975). 
Whether  such  a distance  would  result  in  significantly  different 
paleoenvironmental  conditions  than  those  east  of  the  San 
Andreas  Fault  is  impossible  to  resolve.  Pyroclastic  volcanic 
deposition  dominated  in  the  few  basins  in  existence  east  of 
the  San  Andreas  Fault  in  the  Mojave  Desert  and  only  one 
area  has  yielded  fossils.  By  later  Hemingfordian,  this  vol- 
canism  had  subsided  considerably,  but  most  of  the  basins 
east  of  the  San  Andreas  Fault  were  dominated  by  coarse 
clastic  deposition  as  these  volcanics  were  removed  from  areas 
of  higher  topographic  relief.  Fossils  in  these  coarse  sediments 
are  scattered  and  poorly  preserved.  The  earlier  Hemingfor- 
dian was  not  previously  sampled  (or  recognized,  if  sampled). 
The  Boron  Local  Fauna  samples  this  interval  for  the  first 
time. 

Among  those  southern  California  assemblages  referred  to 
the  Arikareean,  the  Boron  Local  Fauna  shows  some  simi- 
larity to  the  Tick  Canyon  Local  Fauna  (Maxson,  1 930;  Jahns, 
1940)  in  sharing  Trogomys.  Merychyus  cf.  M.  minimus  from 
Boron  is  derived  with  respect  to  M.  calaminthus  from  the 
Tick  Canyon  Local  Fauna.  The  camel  referred  to  Miolabis 
californicus  Maxson,  1930,  from  the  Tick  Canyon  Local  Fau- 
na has  unfused  metapodials  and  is  considerably  larger  than 
the  small  camelid  from  Boron  with  unfused  metapodials. 

Merychyus  cf.  M.  minimus  from  Boron  is  also  derived  with 
respect  to  the  M.  calaminthus  from  the  Late  Arikareean  Black 
Butte  Mine  Local  Fauna  in  the  Hector  Formation  (Wood- 
bume  et  al.,  1974)  and  the  Merychyus  cf.  M.  calaminthus 
from  the  Cady  Mountain  Local  Fauna  (Woodbume,  Miller, 
and  Tedford,  1982).  The  fused,  proximal  metatarsal  from 
the  small  camelid  species  from  Boron  is  considerably  larger 
than  the  Michenia  cf.  M.  agatensis  from  the  latest  Arikareean 
Logan  Mine  Local  Fauna  of  the  Hector  Formation. 

The  Boron  Local  Fauna  shows  little  similarity  to  other 
Arikareean  assemblages  in  California  such  as  the  Tecuya 
Local  Fauna  (Stock,  1920,  1932a),  Kew  Quarry  (Stock,  1932b, 
1935;  Wilson,  1934,  1949),  and  other  localities  in  the  upper 
Sespe  Formation  (Wilson,  1949). 

Among  southern  California  fossil  assemblages  referred  to 
the  late  Hemingfordian,  the  Boron  Local  Fauna  closely  re- 
sembles the  poorly  documented  Phillips  Ranch  Local  Fauna 
(localities  LACM(CIT)  305,  UCMP  V2577)  in  the  Kinnick 
Formation  (or  the  basal  Bopesta  Formation,  James  Quinn, 
personal  communication).  Buwalda  (1916)  and  Buwalda  and 
Lewis  (1955)  recognized  Merychippus  tehachapiensis  Bu- 
walda and  Lewis,  1955  (=  M.  carrizoensis  Dougherty,  1 940— 
see  Munthe,  1 979),  small  and  large  camelids,  Moropus  Marsh, 


Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna  31 


1877,  Merycodus  Leidy,  1854,  canids,  and  felids  from  this 
local  fauna.  Although  not  published,  the  Phillips  Ranch  Local 
Fauna  also  contains  an  oreodont,  peccary,  and  palaeomer- 
ycid.  The  oreodont  is  a small,  high-crowned  species  probably 
referable  to  Merychyus.  It  is  slightly  larger  than  the  Boron 
oreodonts  but  otherwise  similar  to  M.  cf.  M.  minimus.  The 
palaeomerycid  (represented  only  by  upper  teeth)  is  a small, 
high-crowned  species  similar  in  size  and  morphology  to  A. 
occidentalis  or  a small  species  of  Rakomeryx.  The  small  cam- 
el from  Phillips  Ranch  is  intermediate  in  size  between  the 
cf.  Hesperocamelus  sp.  and  the  small  camel  species  from 
Boron  but  appears  to  have  fused  metatarsals  like  the  cf.  Hes- 
perocamelus sp.  from  Boron. 

The  oreodont  from  the  Late  Hemingfordian  Red  Division 
Local  Fauna  in  the  type  section  of  the  Barstow  Formation 
in  the  Mud  Hills  (Woodbume  and  Tedford,  1982),  Meryc- 
hyus ( Metoreodon ) relictus  fletcheri  Schultz  and  Falkenbach, 
1 947,  is  based  on  a heavily  worn,  poorly  preserved  specimen. 
About  all  that  can  be  said  is  that  this  individual  is  larger  than 
the  oreodonts  from  Boron.  The  Red  Division  Local  Fauna 
also  produces  Merychippus  carrizoensis  which  is  absent  at 
Boron. 

Among  other  late  Hemingfordian  fossil  assemblages  in 
southern  California,  the  Boron  Local  Fauna  shares  no  taxa 
with  the  Alvord  Mountain  faunas  from  the  lower  member 
of  the  Barstow  Formation  (localities  D“319,”  D321)  (Byers, 
1960;  Lewis,  1968),  the  Yermo  Hills  Local  Fauna  from  sed- 
imentary rocks  in  the  Yermo  Hills  also  referred  to  the  Bar- 
stow Formation  (Woodbume,  Miller,  and  Tedford,  1982), 
the  fauna  of  the  basal  Punchbowl  Formation  in  the  Cajon 
Valley  (Woodbume  and  Golz,  1972),  Hidden  Treasure  Spring 
Local  Fauna  of  the  Caliente  Formation  in  the  upper  Cuyama 
Valley  (Dougherty,  1940b;  Repenning  and  Vedder,  1961; 
James,  1 963),  the  Vedder  Local  Fauna  of  the  Branch  Canyon 
Formation  (Hutchison  and  Lindsay,  1974;  Lindsay,  1974; 
Munthe,  1979)  or  the  Upper  Cady  Mountain  Local  Fauna 
in  the  northern  Cady  Mountains  (Miller,  1980;  Woodbume, 
Miller,  and  Tedford,  1982).  All  these  have  been  referred  to 
the  late  Hemingfordian  and  all  contain  the  small  Merychip- 
pus carrizoensis,  middle  sized  oreodonts  (except  the  Cady 
Mountain  Local  Fauna)  referred  either  to  Ticholeptus  cali- 
montanus,  Merychyus  ( Metoreodon ) relictus  or  Brachyus  bu- 
waldi,  and  small  camelids  (except  the  Alvord  Mountain  area), 
often  referred  to  Protolabinae.  Although  the  taxonomic  as- 
signments may  be  erroneous,  all  these  oreodonts  are  derived 
with  respect  to  those  from  Boron.  The  only  palaeomerycids 
in  any  of  these  faunas  are  brachyodont  dromomerycines. 

The  Vedder  Local  Fauna  is  the  only  one  of  these  late  Hem- 
ingfordian assemblages  to  also  contain  microvertebrates  ex- 
cept for  5 isolated  specimens  of  Proheteromys  sulculus  from 
the  Cady  Mountain  Local  Fauna.  The  Vedder  Local  Fauna 
shares  only  one  taxon  ( Mookomys ) at  the  generic  level  with 
the  Boron  Local  Fauna,  but  the  Boron  species  is  distinctly 
smaller.  The  Boron  Local  Fauna  lacks  the  squirrels,  eomyids, 
and  Proheteromys  species  found  in  the  Vedder  Local  Fauna, 
and  the  P.  sulculus  found  in  the  Cady  Mountain  Local  Fauna. 
On  the  other  hand,  the  Vedder  Local  Fauna  lacks  Trogomys 
and  Cupidinimus  of  the  Boron  Local  Fauna,  even  though  the 


latter  is  a common  element  in  earliest  Barstovian  and  later 
Miocene  faunas.  The  Vedder  Local  Fauna  is  from  near  shore 
or  possibly  even  marine  rocks,  and  perhaps  the  absence  of 
Cupidinimus  is  environmentally  controlled. 

Taken  on  balance,  the  Boron  Local  Fauna  is  best  consid- 
ered early  Hemingfordian,  a view  consistent  with  the  avail- 
able radiometric  data  and  biostratigraphic  correlations  with 
the  reference  fossil  assemblages  in  Southern  California  and 
the  central  United  States. 

CONCLUSIONS 

Based  on  the  above  discussions,  the  Boron  Local  Fauna: 

1 . is  the  first  diverse  early  Hemingfordian  fossil  vertebrate 
assemblage  from  southern  California, 

2.  is  the  only  diverse  Hemingfordian  microvertebrate  as- 
semblage in  southern  California  east  of  the  San  Andreas 
Fault, 

3.  contains  two  new  mammalian  species,  Cupidinimus  bo- 
ronensis  n.  sp.  (Mammalia,  Rodentia,  Heteromyidae) 
and  Aletomeryx  occidentalis  n.  sp.  (Mammalia,  Artio- 
dactyla,  Palaeomerycidae), 

4.  contains  one  of  the  earliest  North  American  records  of 
the  Family  Colubridae  (Reptilia,  Squamata)  and  the  first 
Hemingfordian  record  west  of  the  Rocky  Mountains, 

5.  contains  the  earliest  species  of  the  genus  Cupidinimus, 

6.  contains  the  second  occurrence  of  Trogomys  (Mam- 
malia, Rodentia,  Heteromyidae),  extending  the  range  of 
this  genus  from  the  Arikareean  into  the  Hemingfordian, 

7.  contains  the  first  association  of  Cupidinimus  and  Tro- 
gomys, the  assumed  ancestor  of  Cupidinimus, 

8.  contains  the  first  Hemingfordian  record  of  a Great  Plains 
species  of  merychyine  oreodont,  Merychyus  minimus 
(Mammalia,  Artiodactyla,  Merycoidodontidae),  west  of 
the  Rocky  Mountains, 

9.  contains  the  oldest  record  of  the  genus  Hesperocamelus 
(Mammalia,  Artiodactyla,  Camelidae), 

10.  contains  the  first  aletomerycine  palaeomerycid,  Aleto- 
meryx occidentalis,  west  of  the  Rocky  Mountains, 

1 1 . provides  a correlative  link  to  the  classical  early  Hem- 
ingfordian assemblages  of  the  mid-continent  because  of 
the  occurrence  of  Merychyus  cf.  M.  minimus  and  Ale- 
tomeryx (both  common  mid-continent  taxa),  and 

12.  provides  a minimum  age  of  Early  Miocene  for  the  Tro- 
pico  Group. 

ACKNOWLEDGMENTS 

I would  like  to  thank  the  geology  staff  of  the  United  States 
Borax  and  Chemical  Corporation,  particularly  Robert  Kis- 
tler,  Sig  Muessig,  James  Minette,  and  Ralph  Barnard  who 
provided  assistance  in  the  collection  of  the  vertebrate  fossils 
and  valuable  insight  into  the  deposits  at  Boron.  I am  indebted 
to  Richard  H.  Tedford  whose  guidance  and  encouragement 
enhanced  the  original  study  of  this  fauna  in  1964-1965.  Rob- 
ert Reynolds  volunteered  considerable  time  in  preparing  some 
of  the  specimens.  Comparative  collections  at  the  American 


32  Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna 


Museum  of  Natural  History  and  the  University  of  California 
at  Berkeley  were  made  available  through  assistance  from 
Richard  H.  Tedford  and  J.  Howard  Hutchison,  respectively. 
Discussions  with  Bruce  Lander  aided  the  analysis  of  the  ore- 
odonts.  John  M.  Harris,  William  A.  Akersten,  and  Lawrence 
G.  Barnes  have  provided  valuable  review  comments.  The 
illustrations  of  artiodactyls  are  by  Karoly  Fogassy  of  the  Uni- 
versity of  California  at  Riverside.  Drafting  of  Figures  1,  2, 
and  60  is  by  Mary  Butler.  Partial  support  was  provided  by 
the  Hewett  Fund  of  the  Department  of  Geological  Sciences, 
University  of  California  at  Riverside. 

LITERATURE  CITED 

Armstrong,  R.L.,  and  R.E.  Higgens.  1973.  K-Ar  dating  of 
the  beginning  of  Tertiary  volcanism  in  the  Mojave  Des- 
ert, California.  Bulletin,  Geological  Society  of  America 
84:1095-1100. 

Auffenberg,  W.  1963.  The  fossil  snakes  of  Florida.  Tulane 
Studies  in  Zoology  10(3):  13 1-2 16. 

Barnard,  R.M.,  and  R.B.  Kistler.  1966.  Stratigraphic  and 
structural  evolution  of  the  Kramer  sodium  borate  ore 
body,  Boron,  California.  In  J.L.  Rau,  Editor.  Second 
Symposium  on  Salt.  Northern  Ohio  Geological  Society 
1:133-150. 

Baskin,  J.  A.  1979.  Small  mammals  of  the  Hemphillian  age 
White  Cone  local  fauna,  northeastern  Arizona.  Journal 
of  Paleontology  53(3):695-708. 

Benda,  W.K.,  R.C.  Erd,  and  W.C.  Smith.  1960.  Core  logs 
from  five  holes  near  Kramer,  in  the  Mojave  Desert, 
California.  United  States  Geological  Survey  Bulletin 
1045F:  1-22. 

Black,  C.C.  1 963.  A review  of  the  North  American  Tertiary 
Sciuridae.  Bulletin,  Museum  of  Comparative  Zoology, 
Harvard  University  1 30(3):  1 13-248. 

Bowen,  C.E.  1954.  Geology  and  mineral  resources  of  the 
Barstow  quadrangle,  California.  California  Division 
Mines  Bulletin  165:7-185. 

Bowser,  C.J.  1965.  Geochemistry  and  petrology  of  the  so- 
dium borates  in  the  non-marine  evaporate  environment. 
Unpublished  Master’s  Thesis,  University  California  at 
Los  Angeles. 

Brattstrom,  B.H.  1958.  New  records  of  Cenozoic  amphib- 
ians and  reptiles  from  California.  Bulletin,  Southern  Cal- 
ifornia Academy  of  Sciences  5 7(1): 5- 12. 

Buwalda,  J.P.  1916.  New  mammalian  faunas  from  Mio- 
cene sediments  near  Tehachapi  Pass  in  the  southern  Sier- 
ra Nevada.  University  of  California  Department  of  Geo- 
logical Sciences  Bulletin  10:78-85. 

Buwalda,  J.P.,  and  G.E.  Lewis.  1955.  A new  species  of 
Merychippus.  Geological  Society  of  America  Profes- 
sional Paper  264G:  147-1 52. 

Byers,  F.M.,  Jr.  1960.  Geology  of  the  Alvord  Mountain 
Quadrangle,  San  Bernardino  County,  California.  United 
States  Geological  Survey  Bulletin  1089-A:1-71. 

Cook,  H.J.  1933.  Faunal  lists  of  the  Tertiary  Vertebrata  of 
Nebraska  and  adjacent  areas.  Nebraska  Geological  Sur- 
vey Paper  5:1-58. 


. 1965.  Runningwater  Formation,  Middle  Miocene 

of  Nebraska.  American  Museum  of  Natural  History 
Novitates  2227:1-8. 

Cope,  E.D.  1873.  Synopsis  of  new  vertebrates  from  Col- 
orado obtained  during  the  summer  of  1873.  Miscella- 
neous Publications,  United  States  Geological  Survey  of 
the  Territories  1:1-19. 

. 1878.  Descriptions  of  new  extinct  Vertebrata  from 

the  Upper  Tertiary  and  Dakota  formations.  Bulletin, 
United  States  Geological  and  Geographic  Surveys  of  the 
Territories  IV:379-396. 

. 1883.  Notes  on  the  geographical  distribution  of 

Batrachia  and  Reptilia  in  western  North  America.  Pro- 
ceedings of  the  Academy  of  Natural  Sciences  of  Phila- 
delphia 1883:10-35. 

Crowell,  J.C.  1975.  The  San  Andreas  Fault  in  southern 
California.  In  J.C.  Crowell,  Editor.  San  Andreas  Fault 
in  southern  California,  a guide  to  the  San  Andreas  Fault 
from  Mexico  to  the  Carrizo  Plain.  California  Division 
Mines  Special  Report  1 18:7-27. 

Dalrhymple,  G.B.  1979.  Critical  tables  for  conversion  of 
K-Ar  ages  from  old  to  new  constants.  Geology  7(11): 
558-560. 

Davidson,  P.W.  1923.  Alticamelus  alexandrae,  anew  cam- 
el from  the  Barstow  Upper  Miocene  of  the  Mojave  Des- 
ert. University  of  California  Publications,  Bulletin  De- 
partment of  Geological  Sciences  14:397-408. 

Dawson,  M.R.  1958.  Later  Tertiary  Leporidae  of  North 
America.  University  of  Kansas  Paleontological  Contri- 
butions 6: 1-75. 

Dibblee,  T.W.,  Jr.  1952.  Geology  of  the  Saltdale  Quadran- 
gle, California.  California  Division  Mines  Bulletin  160: 
1-66. 

. 1958a.  Tertiary  stratigraphic  units  of  the  western 

Mojave  Desert,  California.  Bulletin,  American  Associ- 
ation Petroleum  Geologists  42(1):  135-1 44. 

. 1958b.  Geologic  map  of  the  Boron  Quadrangle, 

Kern  and  San  Bernardino  Counties,  California.  United 
States  Geological  Survey,  Mineral  Investigations  Field 
Studies  Map  MF-204. 

. 1 967.  Areal  geology  of  the  western  Mojave  Desert, 

California.  United  States  Geological  Survey  Professional 
Paper  522:1-153. 

Dickey,  D.D.  1957.  Core  logs  from  two  test  holes  near  Four 
Comers,  San  Bernardino  County,  California.  United 
States  Geological  Survey  Bulletin  1045B:63-79. 

Dorr,  J. A.  1956.  Anceney  local  mammal  fauna,  latest  Mio- 
cene, Madison  Valley  Formation,  Montana.  Journal  of 
Paleontology  30(  1 ):62— 74. 

Dougherty,  J.F.  1940a.  Skull  and  skeletal  remains  of  the 
camel  Paratylopus  cameloides  (Wortman)  from  the  John 
Day  deposits,  Oregon.  Carnegie  Institution  of  Washing- 
ton Publications,  Contributions  in  Paleontology  5 1 4:49- 
58. 

. 1940b.  A new  Miocene  mammalian  fauna  from 

Caliente  Mountain,  California.  Carnegie  Institution  of 
Washington  Publications  514:109-143. 


Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna  33 


Douglass,  E.  1 909.  Dromomeryx,  a new  genus  of  American 
ruminants.  Annals  Carnegie  Museum  V:457-479. 

Downs,  T.  1956.  The  Mascall  fauna  from  the  Miocene  of 
Oregon.  University  of  California  Publications,  Depart- 
ment of  Geological  Sciences  3 1(5):  1 99—354. 

Evemden,  J.F.,  D.E.  Savage,  G.H.  Curtis,  and  G.T.  James. 
1964.  Potassium-argon  dates  and  the  Cenozoic  mam- 
malian chronology  of  North  America.  American  Journal 
of  Science  262:145-198. 

Frick,  C.  1937.  Homed  ruminants  of  North  America.  Bul- 
letin, American  Museum  of  Natural  History  69:1-669. 

Frick,  C.,  and  B.E.  Taylor.  1968.  A generic  review  of  the 
stenomyline  camels.  American  Museum  of  Natural  His- 
tory Novitates  2353: 1-5 1 . 

. 1971.  Michenia,  a new  protolabine  (Mammalia, 

Camelidae)  and  a brief  review  of  the  early  taxonomic 
history  of  the  genus  Protolabis.  American  Museum  of 
Natural  History  Novitates  2444:1-24. 

Galbreath,  E.C.  1953.  A contribution  to  the  Tertiary  ge- 
ology and  paleontology  of  northeastern  Colorado.  Uni- 
versity of  Kansas  Paleontological  Contributions,  Ver- 
tebrata  5:1-120. 

Gale,  H.S.  1946.  Geology  of  the  Kramer  borate  district, 
Kern  County,  California.  California  Journal  of  Mines 
and  Geology  42(4):325-379. 

Galusha,  T.  1975.  Stratigraphy  of  the  Box  Butte  Formation, 
Nebraska.  Bulletin,  American  Museum  of  Natural  His- 
tory 1 56(  1 ):  1 —68. 

Gazin,  C.L.  1930.  A Tertiary  vertebrate  fauna  from  the 
upper  Cuyama  drainage  basin,  California.  Carnegie  In- 
stitution of  Washington  Publications,  Contributions  in 
Paleontology  404:55-76. 

Hall,  E.R.  1930.  Rodents  and  lagomorphs  from  the  later 
Tertiary  of  Fish  Lake  Valley,  Nevada.  University  of  Cal- 
ifornia Publications,  Bulletin  Department  of  Geological 
Sciences  1 9(  1 2):295— 3 1 2. 

Hamilton,  W.R.  1978a.  Cervidae  and  Palaeomerycidae.  In 
VJ.  Maglio  and  H.B.S.  Cooke,  Editors.  Evolution  of 
African  Mammals.  Harvard  University  Press. 

— - — . 1978b.  Fossil  giraffes  from  the  Miocene  of  Africa 
and  a revision  of  the  phylogeny  of  the  Giraffoidea.  Phil- 
osophical Transactions,  Royal  Society  of  London,  Bio- 
logical Sciences  283(999):  165-229. 

Harkson,  J.C.,  and  J.R.  Macdonald.  1967.  Miocene  Bates- 
land  Formation  named  in  southwestern  South  Dakota. 
South  Dakota  Geological  Survey,  Report  of  Investiga- 
tion 96: 1-10. 

Hay,  O.P.  1899.  On  the  names  of  certain  North  American 
fossil  vertebrates.  Science  9(2):593-594. 

Hecht,  M.K.  1956.  A new  xantusiid  lizard  from  the  Eocene 
of  Wyoming.  American  Museum  of  Natural  History 
Novitates  1 174:1-8. 

Hershey,  H.O.  1 902.  Some  Tertiary  formations  of  southern 
California.  American  Geologist  29:349-372. 

Holman,  J.  A.  1970.  Herpetofauna  of  the  Wood  Mountain 
Formation  (Upper  Miocene)  of  Saskatchewan.  Canadian 
Journal  of  Earth  Science  7(5):  13 17-1325. 


. 1973.  Reptiles  of  the  Egelhoff  local  fauna  (Upper 

Miocene)  of  Nebraska.  Contributions,  University  of 
Michigan  Museum  of  Paleontology  24(1 2):  125-1 34. 

. 1975.  Herpetofauna  of  the  Wakeeney  local  fauna 

(Lower  Pliocene:  Clarendonian)  of  Trego  County,  Kan- 
sas. In  G.R.  Smith,  and  N.E.  Friedland,  Editors.  Studies 
on  Cenozoic  Paleontology  and  Stratigraphy.  University 
of  Michigan,  Museum  of  Paleontology:49-74. 

. 1979.  A review  of  North  American  Tertiary  snakes. 

Publications  of  the  Museum,  Michigan  State  University, 
Paleontological  Series  1 (6):203— 260. 

Honey,  J.G.,  and  B.E.  Taylor.  1978.  A generic  revision  of 
the  Protolabidini  (Mammalia,  Camelidae),  with  a de- 
scription of  two  new  protolabidines.  Bulletin,  American 
Museum  of  Natural  History  161  (3):369— 425. 

Hutchison,  J.H.,  and  E.H.  Lindsay.  1974.  The  Heming- 
fordian  mammal  fauna  of  the  Vedder  locality,  Branch 
Canyon  Formation,  Santa  Barbara  County,  California. 
Part  1:  Insectivora,  Chiroptera,  Lagomorpha  and  Ro- 
dentia  (Sciuridae).  PaleoBios  15:1-19. 

Jahns,  R.H.  1940.  Stratigraphy  of  the  easternmost  Ventura 
Basin,  California,  with  a description  of  a new  Lower 
Miocene  mammalian  fauna  from  the  Tick  Canyon  For- 
mation. Carnegie  Institution  of  Washington  Publica- 
tions, Contributions  in  Paleontology  514:145-194. 

James,  G.T.  1963.  Paleontology  and  nonmarine  stratigra- 
phy of  the  Cuyama  Valley  Badlands,  California.  Part  1. 
Geology,  faunal  interpretations  and  systematic  descrip- 
tions of  Chiroptera,  Insectivora  and  Rodentia.  Univer- 
sity of  California  Publications  Geological  Sciences  45: 
1-162. 

Kellogg,  L.  1910.  Rodent  fauna  of  the  late  Tertiary  beds 
of  Virgin  Valley  and  Thousand  Creek,  Nevada.  Uni- 
versity of  California  Publications,  Bulletin  Department 
of  Geological  Sciences  5:41  1-437. 

Klininger,  D.  1968.  Rodents  of  the  Mio-Pliocene  Norden 
Bridge  local  fauna,  Nebraska.  The  American  Midland 
Naturalist  80(l):65-74. 

Korth,  W.W.  1979.  Geomyoid  rodents  from  the  Valentine 
Formation  of  Knox  County,  Nebraska.  Annals  Carnegie 
Museum  48(16):287-3 10. 

Leidy,  J.  1858.  Notice  of  remains  of  extinct  Vertebrata  from 
the  valley  of  the  Niobrara  River,  collected  during  the 
exploring  expedition  of  1857,  in  Nebraska.  Proceedings 
of  the  Academy  of  Natural  Sciences  of  Philadelphia  10: 
20-29. 

Lewis,  G.E.  1955.  A new  species  of  Merychippus.  Geolog- 
ical Society  Professional  Paper  364-G:  147-1 51. 

. 1968.  Stratigraphic  paleontology  of  the  Barstow 

Formation  in  the  Alvord  Mountain  area,  San  Bernar- 
dino County,  California.  United  States  Geological  Sur- 
vey Professional  Paper  600-C:C75-C79. 

Lindsay,  E.H.  1972.  Small  mammal  fossils  from  the  Bar- 
stow Formation,  California.  University  of  California 
Publications  Geological  Sciences  93:1-104. 

. 1974.  The  Hemingfordian  mammal  fauna  of  the 

Vedder  locality,  Branch  Canyon  Formation,  Santa  Bar- 


34  Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna 


bara  County,  California.  Part  II.  Rodentia  (Eomyidae 
and  Heteromyidae).  PaleoBios  16:1-20. 

Loomis,  D.P.,  B.F.  Cox,  and  D.P.  Whistler.  1983.  Basin 
analysis  of  the  Ricardo  Formation  and  implications  for 
Miocene  history  of  the  Garlock  Fault.  Geological  Society 
America,  Abstract,  Cordilleran  Meeting,  May,  1983. 

Loomis,  F.B.  1924.  Miocene  oreodonts  in  North  America. 
Bulletin,  American  Museum  of  Natural  History  51(1): 
1-37. 

Lull,  R.S.  1 920.  New  Tertiary  artiodactyls.  American  Jour- 
nal of  Science  50:83-130. 

. 1921.  New  camels  in  the  Marsh  collection.  Amer- 
ican Journal  of  Science  50:392-404. 

Macdonald,  J.R.  1949.  A new  Clarendonian  fauna  from 
northeastern  Nevada.  University  of  California  Publi- 
cations, Geological  Sciences  28(7):  173-1 94. 

. 1956.  A new  Clarendonian  Mammalian  Fauna  from 

the  Truckee  Formation  of  western  Nevada.  Journal  of 
Paleontology  30(1):  186-202. 

. 1963.  The  Miocene  faunas  from  Wounded  Knee 

area  of  western  South  Dakota.  Bulletin,  American  Mu- 
seum of  Natural  History  125:139-238. 

. 1970.  Review  of  the  Miocene  Wounded  Knee  fau- 
nas of  southwestern  South  Dakota.  Bulletin,  Los  Angeles 
County  Museum  of  Natural  History,  Science  8:1-82. 

Macdonald,  J.R.,  and  J.C.  Harksen.  1968.  Rosebud  For- 
mation in  South  Dakota.  South  Dakota  Geological  Sur- 
vey, Report  of  Investigations  97:1-13. 

Martin,  J.E.  1976.  Small  mammals  from  the  Miocene 
Batesland  Formation  of  South  Dakota.  Contributions  to 
Geology,  University  of  Wyoming  14(2):69-98. 

Matthew,  W.D.  1901.  Fossil  mammals  of  the  Tertiary  of 
northeastern  Colorado.  Memoirs,  American  Museum  of 
Natural  History  1:355-448. 

. 1904.  Notice  of  two  new  Oligocene  camels.  Bulletin 

of  the  American  Museum  of  Natural  History  20:211- 
215. 

. 1924.  Third  contribution  to  the  Snake  Creek  fauna. 

Bulletin  American  Museum  of  Natural  History  50(2): 
59-210. 

Matthew,  W.D.,  and  H.J.  Cook.  1909.  A Pliocene  fauna 
from  western  Nebraska.  Bulletin  American  Museum  of 
Natural  History  26:361-414. 

Matthew,  W.D.,  and  J.R.  Macdonald.  1960.  Two  new 
species  of  Oxydactylus  from  the  middle  Miocene  Rose- 
bud formation  in  western  South  Dakota.  American  Mu- 
seum of  Natural  History  Novitates  2003:1-7. 

Maxson,  J.H.  1930.  A Tertiary  mammalian  fauna  from  the 
Mint  Canyon  Formation  of  southern  California.  Car- 
negie Institute  of  Washington  Publications,  Contribu- 
tions in  Paleontology  404:77-1 12. 

McKenna,  M.C.  1965.  Stratigraphic  nomenclature  of  the 
Miocene  Hemingford  Group,  Nebraska.  American  Mu- 
seum of  Natural  History  Novitates  2228:1-21. 

Merriam,  J.C.  1911.  The  Tertiary  mammal  beds  of  Virgin 
Valley  and  Thousand  Creek  in  northwestern  Nevada; 
part  II,  vertebrate  fauna.  University  of  California  Pub- 


lications, Bulletin  Department  of  Geological  Sciences  6: 
199-304. 

. 1919.  Tertiary  mammalian  faunas  of  the  Mojave 

Desert.  Bulletin,  University  of  California  Department 
of  Geological  Sciences  1 1:437-585. 

Miller,  S.T.  1980.  Geology  and  mammalian  biostratigra- 
phy of  a part  of  the  northern  Cady  Mountains,  Califor- 
nia. United  States  Geological  Survey  Open  File  Report 
80-978:1-121. 

Munthe,  J.  1979.  The  Hemingfordian  mammal  fauna  of 
the  Vedder  locality.  Branch  Canyon  Sandstone,  Santa 
Barbara  County,  California.  Part  III:  Carnivora,  Peris- 
sodactyla,  Artiodactyla  and  summary.  PaleoBios  29:1- 
22. 

Peterson,  O.A.  1904.  Osteology  of  Oxydactylus.  Annals 
Carnegie  Museum  2(3):434-475. 

. 1906.  The  Miocene  beds  of  western  Nebraska  and 

eastern  Wyoming  and  their  vertebrate  fauna.  Annals  of 
the  Carnegie  Museum  IV:21-72. 

. 1911.  A new  camel  from  the  Miocene  of  western 

Nebraska.  Annals  of  the  Carnegie  Museum  7(2):260- 
266. 

Reeder,  W.G.  1956.  A review  of  Tertiary  rodents  of  the 
Family  Heteromyidae.  Unpublished  Ph.D.  Dissertation, 
University  of  Michigan. 

. 1960.  A new  rodent  genus  (Family  Heteromyidae) 

from  the  Tick  Canyon  Formation  of  California.  Bulletin 
Southern  California  Academy  Sciences  59(3):  12 1-1 32. 

Repenning,  C.A.,  and  J.G.  Vedder.  1961.  Continental  ver- 
tebrates and  their  stratigraphic  correlation  with  marine 
mollusks,  eastern  Caliente  Range,  California.  United 
States  Geological  Survey  Professional  Paper  424:C235- 
C239. 

Schatzinger,  R.A.  1980.  New  species  of  Palaeoxantusia 
(Reptilia:  Sauria)  from  the  Unitan  (Eocene)  of  San  Diego 
County,  California.  Journal  of  Paleontology  54(2):460- 
471. 

Schultz,  C.B.  1938.  The  Miocene  of  western  Nebraska. 
American  Journal  Science  35:441-444. 

Schultz,  C.B.,  and  C.R.  Falkenbach.  1940.  Merycochoeri- 
nae,  a new  subfamily  of  oreodonts.  Bulletin  Museum  of 
Natural  History  77(5):2 13-306. 

. 1941.  Ticholeptinae,  a new  subfamily  of  oreodonts. 

Bulletin  American  Museum  of  Natural  History  79(  1 ):  1 — 
105. 

. 1947.  Merychyinae,  a subfamily  of  oreodonts.  Bul- 
letin, American  Museum  of  Natural  History  88(4):  161— 
285. 

Shotwell,  J.A.  1967.  Later  Tertiary  geomyoid  rodents  of 
Oregon.  University  of  Oregon,  Museum  of  Natural  His- 
tory Bulletin  9:1-51. 

Sinclair,  W.J.  1915.  Additions  to  the  fauna  of  the  Lower 
Pliocene  Snake  Creek  beds  (results  of  the  Princeton  Uni- 
versity 1914  expedition  to  Nebraska).  Proceedings, 
American  Philosophical  Society  54:73-95. 

Skinner,  M.F.,  S.M.  Skinner,  and  R.J.  Gooris.  1977.  Stra- 
tigraphy and  biostratigraphy  of  late  Cenozoic  deposits 


Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna  35 


in  central  Sioux  County,  western  Nebraska.  Bulletin, 
American  Museum  of  Natural  History  1 58(5):265— 37 1 . 

Stevens,  M.S.  1977.  Further  study  of  Castolon  local  fauna 
(Early  Miocene),  Big  Bend  National  Park,  Texas.  Pearce- 
Sellards  Series,  Texas  Memorial  Museum  28:1-69. 

Stevens,  M.S. , J.B.  Stevens,  and  M.R.  Dawson.  1969.  New 
Early  Miocene  formation  and  vertebrate  local  fauna,  Big 
Bend  National  Park,  Brewster  County,  Texas.  Pearce- 
Sellards  Series,  Texas  Memorial  Museum  15:1-53. 

Stirton,  R.A.  1944.  Comments  on  the  relationships  of  the 
cervoid  Family  Palaeomerycidae.  American  Journal  Sci- 
ence 242:633-655. 

Stock,  C.  1920.  An  early  Tertiary  vertebrate  fauna  from 
southern  Coast  Ranges  of  California.  Bulletin,  Depart- 
ment Geology,  University  of  California  12:267-276. 

. 1932a.  Additions  to  the  mammalian  fauna  from 

the  Tecuya  beds,  California.  Carnegie  Institution  Wash- 
ington 418:87-93. 

. 1932b.  An  Upper  Oligocene  mammalian  fauna  from 

southern  California.  Proceedings,  National  Academy  of 
Sciences  18:550-554. 

. 1935.  Artiodactyla  from  the  Sespe  of  the  Las  Posas 

Hills,  California.  Carnegie  Institution  Washington  Pub- 
lications, Contributions  in  Paleontology  453:1  19-125. 

Storer,  J.A.  1970.  New  rodents  and  lagomorphs  from  the 
Upper  Miocene  Wood  Mountain  Formation  of  southern 
Saskatchewan.  Canadian  Journal  of  Earth  Sciences  7(4): 
1125-1129. 

. 1975.  Tertiary  mammals  of  Sasakatchewan,  Part 

III:  the  Miocene  fauna.  Royal  Ontario  Museum,  Life 
Sciences  Contributions  103:1-134. 

Sutton,  J.F.  1977.  Mammals  of  the  Anceney  local  fauna 
(Late  Miocene)  of  Montana.  Unpublished  Ph.D.  Dis- 
sertation, Texas  Tech  University. 

Taylor,  B.E.,  and  S.D.  Webb.  1976.  Miocene  Leptomeryc- 
idae  (Artiodactyla,  Ruminantia)  and  their  relationships. 
American  Museum  of  Natural  History  Novitates:  2566: 
1-22. 

Tedford,  R.H.,  T.  Galusha,  M.F.  Skinner,  B.E.  Taylor,  R. 
Fields,  J.R.  Macdonald,  T.H.  Patton,  J.M.  Rensberger, 
and  D.P.  Whistler.  1973.  Faunal  succession  and  bio- 
chronology of  the  Arikareean  through  Clarendonian  in- 
terval (Miocene  Epoch),  North  America.  Geological  So- 
ciety of  America,  Abstracts  with  Programs  5(7):837- 
838. 

Thorpe,  M.R.  1937.  The  Merycoidodontidae,  an  extinct 
group  of  ruminant  mammals.  Memoirs,  Peabody  Mu- 
seum of  Natural  History  111(4):  1—428. 

Turner,  D.L.  1970.  Potassium-argon  dating  of  Pacific  Coast 
Miocene  foraminiferal  stages.  Geological  Society  of 
America  Special  Paper  124:91-129. 

Webb,  S.D.  1965.  The  osteology  of  Camelops.  Bulletin, 
Los  Angeles  County  Museum  of  Natural  History  1:1- 
54. 

Webb,  S.D.,  and  B.E.  Taylor.  1980.  The  phylogeny  of  horn- 
less ruminants  and  a description  of  the  cranium  of  Ar- 
chaeomeryx.  Bulletin,  American  Museum  of  Natural 
History  167(3):  12 1-1 57. 


Wilson,  R.W.  1934.  Two  rodents  and  a lagomorph  from 
the  Sespe  of  the  Las  Posas  Hills,  California.  Carnegie 
Institution  Washington  Publications,  Contributions  in 
Paleontology  453: 1 1-17. 

. 1939.  Rodents  and  lagomorphs  of  the  Late  Tertiary 

Avawatz  fauna,  California.  Carnegie  Institution  Wash- 
ington Publications,  Contributions  in  Paleontology  514: 
31-38. 

. 1 949.  Rodents  and  lagomorphs  of  the  upper  Sespe. 

Carnegie  Institution  Washington  Publications,  Contri- 
butions in  Paleontology  584:51-65. 

. 1960.  Early  Miocene  rodents  and  insectivores  from 

northeastern  Colorado.  University  of  Kansas  Paleon- 
tological Contributions,  Vertebrata  7:1-92. 

Wood,  A.E.  1931.  Phylogeny  of  the  heteromyid  rodents. 
American  Museum  Novitates  501:1-19. 

. 1932.  New  Miocene  rodents  from  the  Miocene  of 

Florida.  Florida  State  Geological  Survey  Bulletin  10:43- 
51. 

. 1935.  Evolution  and  relationship  of  the  heteromyid 

rodents  with  new  forms  from  the  Tertiary  of  western 
North  America.  Annals  Carnegie  Museum  24:73-262. 

. 1936a.  Geomyid  rodents  from  the  middle  Tertiary. 

American  Museum  of  Natural  History  Novitates  866: 
1-23. 

. 1936b.  Fossil  heteromyid  rodents  in  the  collections 

of  the  University  of  California.  American  Journal  of 
Science,  Fifth  Series  32(188):1 12—1 19. 

. 1937.  Additional  material  from  the  Tertiary  of  the 

Cuyama  basin  of  California.  American  Journal  of  Sci- 
ence 33:29-43. 

Woodbume,  M.O.,  and  D.J.  Golz.  1972.  Stratigraphy  of 
the  Punchbowl  Formation,  Cajon  Valley,  southern  Cal- 
ifornia. University  of  California  Publications  Geological 
Sciences  92: 1-57. 

Woodbume,  M.O.,  and  R.H.  Tedford.  1982.  Litho-  and 
biostratigraphy  of  the  Barstow  Formation,  Mojave  Des- 
ert, California.  In  Guidebook,  Geologic  Excursions  in  the 
California  Desert.  Cordilleran  Section,  78th  Annual 
Meeting,  Geological  Society  of  America:47-64. 

Woodbume,  M.O.,  and  D.P.  Whistler.  1973.  An  Early 
Miocene  oreodont  (Merychyinae,  Mammalia)  from  the 
Orocopia  Mountains,  southern  California.  Journal  of 
Paleontology  47(5):908— 9 1 2. 

Woodbume,  M.O.,  S.T.  Miller,  and  R.H.  Tedford.  1982. 
Stratigraphy  and  geochronology  of  Miocene  strata  in  the 
central  Mojave  Desert,  California.  In  Guidebook,  Geo- 
logic Excursions  in  the  California  Desert.  Cordilleran 
Section,  78  th  Annual  Meeting,  Geological  Society  of 
America:47-64. 

Woodbume,  M.O.,  R.H.  Tedford,  M.S.  Stevens,  and  B.E. 
Taylor.  1974.  Early  Miocene  mammalian  faunas,  Mo- 
jave Desert,  California.  Journal  of  Paleontology  48(1): 
6-26. 


Accepted  25  May  1984. 


36  Contributions  in  Science,  Number  355 


Whistler:  Boron  Local  Fauna 


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